The Butterflies of California

T H E B U T T E R F L I E S O F C A L I F O R N I A John F. Emmel, Thomas C. Emmel, and Sterlin g O. Mattoo n Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon DOI:10.5281/zenodo.17313658 Table of C ontents Preface Chapter 1. Early Lepidopter ists in California 1 Chapter 2. The Geological Landscape of California 62 Chapter 3. The Relation o f the California Climate to the Butterfly Fauna 66 Chapter 4. Plant Communities o f California 69 Chapter 5. Patterns o f Geography and Species Diversity in the State o f California 124 Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon Chapter 6. The California Islands and Their Butterfly Faunas 135 Chapter 7. Effects o f Human A ctivities on L epidopteran Populations in California 139 Chapter 8. S pecies Accounts of the Butterflies and Skippers of California 150 Map of localities 752 Literature cited 753 PREFA CE After the p ublicatio n of T he Butterf lies of Sout hern Califo rnia (T. C. Emmel an d J. F. Emmel , 1973), a number of colleagues urged us to prepa re a new book of similar depth covering a ll of the sta te of Califo rnia. The fortunate availab ility of S terling O . Mattoon as a co - author meant that the northern California fauna could be covered wit h the help of an expert resident lepidopterist whose legendary field work in that part of the state would add much to the pr oposed work. Accordingly, the three of us joined forces in 1974 and signed a contract with Stanford University Press, ambitiously estimating that the new book would be published by 1979. Little did we realize the immensity of the task before us in 1974. We began with a tho rough su rvey form that was mailed to all we stern membe rs of the Lepidopterists' Society, requesting information on unusual distributional records, dates, etc., for the Califo rnia fa una as lis ted on a p relimi nary draft of the che cklist o f all recorded butterfly species for the state. That list ra pidly expanded to include dozens of additional species, as we began to rece ive these check lists back from willing collab orators. The help continued to pour in from hundreds of lepidopterists as w e pursued the p reparation of th is book over the ensuing 18 years. Additionally, the lengthy gestation time for this book allowed John F. Emmel in particular to eventually rear and photograph the life history stages of a ll of the spec ies and virtu ally all of the subspecies of butterflies and skippers in the state of C alifornia. Map ping of distrib utiona l records was fa cilitated by visits to al l major priv ate and pu blic colle ctions in the stat e, as well as specia lly targete d field tr ips each year by the th r ee authors and willing colleagues to unknow n or relatively unknown parts of California. In th e course of that effort, several new sp ecies and more than 100 new subspecie s of butterfl ies were des cribed in for mal publica tions, part icular ly in the work entit led Systematics of Western North Ameri can Butterfli es (1992). Thus the present book represents an unusually thorough coverage of an incredibly diverse fauna in a hug e state comprising some of the most var ied geographical an d ecological environments in the northern he misphere. We know there are errors of commission and omission in a work of this size, but we hope that the reader will be encouraged to add to the knowledge represented i n this volume and will communicate new data and discoveries to us so that they may be incorporated in future printi ngs of this book. W e also hope the appearance of this book will generate a great many independent publications on life histories, foodplant records, and all the basic ecology, behavior, genetics, and evolutionary biosystematic work that a basic knowledge of the California fa una should stimulate and encourage. John F. Emmel Thomas C. Emmel Sterling O. Mattoon Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon Chapter 1 EARLY LEPI DOPT ERISTS IN CALI FORNIA Butterflies have probably fascinated mankind as long as both have existed on the earth. Even today, prim itive tribes in areas as far apart as the So uth American Amazon and the interior highlands of New Guinea know inti mately the natural world around them and inclu de butterflies as objects to admire, revere, or even d read, depending on the particular folklore surrounding them. As agricu ltural prac tices started an d centers o f civili zation gr ew, mankin d could no t fail t o note the depredat ions of certain sp ecies on cultiv ated crops, or the mass emergen cies of migran ts like the Painted Lady, or the sheer beauty and fragility of the adult butte rfly. Thus w ith the invention of writing, early chronicles speak of such observations i n Greek and Roman times, and eventually in medieval Europe. But it was not unt il the Age of Exploration, as European adventurers sail ed the oceans of the w o rld and explored the newly opened continents, that the study of natural history started on an org anized, indeed enthusiastically pursued, basis. Collec tions began to be amassed by institutions and individuals alike, both fr om home territories and abroad, during the seventeenth and eighteenth cent ury, and Linnaeus himself laid the groundwork for formal b iological classifi cation, inc luding p ersonally naming butterfl ies even from th e Americas, in his great Systemae Naturae work of 1758. Virtually every new exploring expedition sent back unique and wonderful species of plants and animals to Europe to be named. President Thomas Jefferson started the same tradition in America on 20 June 1803 by charging M eriwether L ewis and Willia m Clark to go forth and exp lore his great ne w Louisia na Purchase and bring back all manner of information on the region's natural histo ry. A year later, they set o ut from St. Louis up the M issouri Rive r. Pushing westw ard from th e headw aters of th e Missouri to the Pacific Ocean it self, the Lewis and Clark Expedit ion was but the fi rst of many government and private expeditions to follow t hroughout the West during the nineteenth century. Among them were lepidopterists traveling with Russian explorers, Army expeditions, gold miners, survey parties, and finally even museums and pr ivate collector sponsors. They set the stage fo r the wealth o f le pidopte rist intere st in Califo rnia's bu tterfly fa una during the twenti eth century . In this first c hapter, the n, we trace th e history of L epidoptera c ollecting a nd butter fly study in California. Not all individuals can be mentioned, of course, but the story covers the highlights -- both chief contributors and interesting oddities -- and gives the reader an appreciation for the sweep of history and the remarkable continuity and depth that Lepidopter a work has enjoyed for the past two centuries in California. The story is st ill being written today by thousands of naturalis ts and scien tist s interes ted in the bu tterflies of thi s great state, and it is ho ped that the readers of this book will see themsel ves as an int egral part of the e ffort to un derstand a little mo re about our natural world -- especially t hat world of th e Californ ia butter fl y fauna. Looking back over the long stretch of years involved in the development of California butterfly study, w e may identify six more- or - less distinct pe riods of act ivity. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 2 1791-1848: The Russians & E arliest Explore rs during the Span ish -Colonial Period in C alifornia. 1849-1859: The Gold Rush Era, Mexican Boundary Sur vey, and Pacific Railroad Survey. 1860: 1870: The State Geologica l Survey Decade. 1870-1890: The Federal Surv eys of the West: Period following t he Opening of the Tran scontinental R ailroad. 1890-1940: The Golden Era of California Lepidoptera Discovery. 1940- Present: Th e Modern Age of Lepidop tera Study: Recent Lep idopterists in California . In the pages that follow, w e shall follow the above outline, discussing the general activities and the principal figures involved during each period. I. THE SP ANISH - COLONIAL PE RIOD (179 1- 1848) Several e arly voy ages of explo ratio n brought th e first n aturalists to coll ect butte rflies in California. These were Russians, surprisingly e nough, who were surveying the Pacific C oast of North Amer ica from Al aska south t o central C alifornia an d collectin g natural history specimens, mostly bo tanical but so me entomol ogical (Gr innell, 1914 ). Among these latter tre asures were some of Ca lifornia' s most notab le butterf lies. Early Russian E xplorers The first n aturalis t to collec t butterfli es and ot her insects in Cali fornia were R ussian scientists and explorers. Grigory I. von Longsdorff, a Russian botanist and zoologist, shipped plant and anima l specimens back to the Russian Academy of Sciences in 1806, but unfortunately these collections were lost and never described. In October, 1816, the Russian navigat or, Captain Otto von Kotzebue brought the nat uralist Adelbert von Chamisso and the entomologist Johann Friedrich Eschsch oltz to San Francisco abo ard the brig Rurik . When they landed on Yerba Buena Island, they found the Chalcedon Checker spot, Euphydryas chalcedona , as verified later by H. H. Behr who saw the specimen in the Berlin Museum. On the same voyage, they visited San Jose and Monterey, and Esc hscholtz collect ed insects in the Fort R oss area on his second trip wi th Kotzedue in 1823 to 1826 (Wells, 1987). The beautiful California Poppy, E schscholtzia californica , was named by Chamisso for his collecting companion Eschscholtz. It is an hist orical oddity that our State Flow er, the California Poppy, and the State Butterfly, the California Dogfa ce ( Zerene eurydice , o riginally Z. wo snesenskii) , w ere named very early fo r the Russians. Johann Friedrich E schscholtz (1793 -1831) Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 3 Though living only 38 years, Eschscholt z was the outstanding entomologist during the Russian occupation of California. He was born in Russia on November 1, 1793 and died at his birthplace on May 7, 1831. Trained in the medical profession, he acted as phy sici an and n aturalist on the ship Rurik which sailed around the world under the command of Otto von Kotzebue during the years 1815 - 1818. H is close friend, a German poet and naturalist, Adelbert von Chamisso, received most of Eschs choltz's captu res except for the insects. This ship a rrived in Sa n Francis co Bay on October 1, 1816, and remained until November 1. He confined his collecting on this trip to the Presidio at San Francisco, the mission and the area in between, and did not visit the Russian colon y at Fort Ross in 1816. Lit tle was taken in the way of insec ts but he did supposedly take two very commo n California s pecies of butte rflies, th e Monarch, Danaus plexippus , and the Chalce don Checkerspot, Euphydryas chalcedona . According to Fordyce G rinnell, Herman H. Behr saw the actual speci mens taken by Eschscho ltz in the co llections of th e Berlin Museum (G rinnell, 1914). It is not difficult to believe that Eschscholtz caught Monarchs in the fall of 1816, but it is less ce rtain how he obtained the normally spring - flying E. chalced ona on the coast. Perhaps the sing le type specimen was from an unknown collector in the Russian colony. ( However, R. L. Langston ( in litt .) has found small numbers of f resh E. chalcedona flyin g in October in 1985 and 1991 in the San Bruno Mountains next to San F rancisco Bay, so a sharp entomologist such as Eschscholtz could have picked up an unusually attractive butterfly like chalcedona if a similar f all brood w as flying in 1816.) Essig ( 1972) believes that these butterfly specimens must have been p laced in the Berlin Museum by Chamisso, whose plant colle ction is there. Incidentally, it was on the 1816 trip that the California Poppy wa s collected, w hich Chamiss o afterwards named E schscholtzia ca lifornica because of his friendship with Eschsc holtz. Following their visit to California, they went to Hawaii in November 1816. He collected on the island of Oahu the beautiful endemic nymphalid which he subsequently named Vanessa tameamea . Eschscholtz made a second voyage with Kotzebue in th e ship Predpriaetie ( Enterprise ) during the years 1823 to 1826. Esc hscholtz was chief naturalist and amassed a ve ry large and important co llection, includin g specimens fro m a Califo rnia visit from Septemb er 27 to N ovember 25, 1824, at San Francisco Bay. However, collections were made throughout the area, to Santa Clara, San Rafael, Fort Ross, Bodega Bay, and t he lower portions of the Sacra mento River, possibly to the present site of Rio Vista. Although he collected many new sp ecies of Coleoptera, Eschscholtz had less luc k with the Lepidoptera, since the ir delayed syste matic study commenced at the time (starting in 1850 or 1851) that Boisduval was already rapidly describ ing the new Californ ia butterfl ies collected by Lorquin. Ilya Vosnesen sky Ilya Gavrilovish Vosnesensky followed the first Russian butterfly collectors by 24 years , when he arriv ed on a special tr ip to the Russian - American colonies at Fort Ross in Sonoma County on July 20, 1840, and remained until Septem ber 3, 1849 (Essig, 1972; Wells, 1987). Only 23 years old in 1839, he was selected to go to the Russian colonies in North Americ a, and his efforts were so productive and successful that he extended his planned three - year journey to ten years. His Fort Ross sojourn w as especially producti ve in insects, plants, and even mammals. Fort R oss had been established in California in 1812 with about 30 Rus sians and 150 Aleuts from Alaska, when Ivan A. Kusk of built a fortifi ed village an d called it " Ross" (a wo rd from the same roo t as "Rus sia" ), as Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 4 headquarters for their farms (t o provide food for the Russian - Alaskan colony) and for trading wit h the Spanish and the Indians. This site was some 13 miles no rth of the presently named Russian River, and 18 miles north of Bodega Bay. Vosnesensky w as quite a successful collector an d assembled a large collection of insects from the San Francisco Bay region, extending from Fort Ross and the Russian River to Bodega Bay and New Helvetia (S utter's Fort). He was the firs t to collect the Cal if ornia Dogfa ce butterfly , Zerene eurydice Boisduval, which was named by the Russian entomologist, Edouard Menetries, as Zerene wosnesenskii Ménétriés in 1855. However, shortly before his name was published, the French entomologist J. H. A. Boisduval publ ished the de scription o f the same b utterf ly (collecte d by Lorquin), giving the name C. eurydice priority. Vosnesensky is perhaps best known in California history f or having been the first person other than native Indians to climb and name Mount St. Helen a (4,343 feet high) on June 12, 1841. This is the most conspicuous mounta in in the vicinity of the Fort Ross s ite. On the summit, he and his Russian companion placed a s mall inscribed copper plate announcing the event. This plate was removed in 1853 and was subs equently destroyed in the San Francisco fire of Apr il 1906. On the 100th anniversary of the founding of Fort Ross in June 1912, the plaque was restored and placed again on the summit. It is probable that a number of our California butterflies would have been named from Vosne sensky's collecting in No rth America, had Ménétriés not been forestalled by Boisduval (1852 and following pa pers) publishing the exciting finds m ade by his collector and friend, Pierre Lorquin, in California just ten years after Vosnesensky's visit there. II. THE GOL D RUSH ERA (1 849 - 1859) Pierre Josep h Michael Lorquin The most famous of our early Calif ornia collectors, Lorquin was born in Valenciennes, France, on July 2, 1797, and died in Par is, France on February 8, 1873. While Essig (1972) summariz es many of the deta ils of Lo rquin's life with a ccuracy , the itiner ary outlined by him has been corrected and given in greater detail in Emmel, Emmel, and Mattoon (1993a), from which the following account is taken. During his 75 years of life, Lorquin explored and collected in a number of areas around the world. As a student at the University of Douay, he became a graduate with honors and became a notary clerk, rising rapidly and taking a position in Algiers in 1848, where he co llected a gre at deal of in teresting lepidoptero logical m aterial. O n hearing of the discovery of gold in California in 1850, he left Algiers and headed for the new El Dorado, not just for gold but for the lepidopterological terra incognita . Upon arrival in California, he found an unexplored field for Lepidoptera and set to work t o supply his intimate friend, the great F rench lepidopterist J. A. Boisduval (family physician for the Lor quins back in France), with specimens. His family followed in 1852 and he remained in the state until 1859. Boisduval (1868) has a fascinating preface to his paper which we quote part of below to give the flavor of what Lorquin faced in the 1849 gold rush: Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 5 "A naturalist known to all for his zeal and love of science, M. Lorquin, wa s seduced as the others have been by the shining mirage and emba rked for California to remake a fortune of which he had been so unjustly stri pped. He left in 1849, his heart full of hope, bel ieving that in this Eldorado one had only to bend over, so to speak, and pick up the gold nuggets. As soon as he landed, and without pausing a moment to rest, he hastened t o the Placers and worked lik e a com mon laborer, with as mu ch cou rage as ene rgy, from time to t ime findin g a few particle s of the prec ious meta l; but the result of this hard work were not even enough to pay his expenses in a land where everything had a fabu lous price: 30 francs for a dozen of eggs, and the rest in proportio n! Not knowing what to do, he braced hims elf against adversity, left the Placers, returned to San Francisco and, putting aside all pride, he did what many others were doing: he practiced many occupations well outside his usual ones, and this permitted him t o live and amass a little money. From this moment dates the honorable reputation he as acquired today, by force of economy, for work and perseverance. Having acquired a little free time, and having hi s family with him (they had com e out to join him), he felt awakening in himself a love of natural science and above all of entomology. To indulge in these interests, he did not hesitate to endure rude privations and much hard work. He first explored all the environs of San Francisco, then the shores of the Sacramento and the Plume [Feather], made voyages into the chain of the Sierra Nevada and ventured even to the forests of the interior, bravi ng the bear' s tooth and the rattlesnake's fang. These dangerous excursions, made in different seasons over two years, fur nished him with fine collections of insects of which he sent us, eighteen years ago, two extremely outstanding shipments. A large part of the Lepidoptera being unknown and entirely new, we published their descriptio ns in the Annals of the Ento mological S ociety of France in t he form of a small loc al fauna. Since that time, M. Lorquin's zeal has not slackened. On the contrary, having more leisure time he undertook voyages to unex plored regions: He went to visit the mountains of the north; penetrated far to the Eas t; and later went into the homeland of the Apaches, as far as Los Angeles in Sonar. T he results of these c ollec ting trips in these d iverse part s of Califo rnia con firmed ful ly to what we would expect of such a man as M. Lorquin. T his indefatigable entomologist sent us three succes sive shipments of speci mens collected in th ese different loc alities, each o ne including many new species, w hose number ex ceeds that of the specie s we made known in the small fauna just mentioned. We believed that with these new materials, w e would render a service to l epidopterists by giving them now a second part, no less interesting than the first. To make this little work more useful and a s complete as po ssible we have, in the intere st of thos e who do not have our earlier work, reproduced all the diagnostic phrases of the species described in the Annals of th e Entomological Society; we also give, in the form of an appendix at the end, a list of the Noctuids and Geometers discovered by M. Lorqui n and co mmunicated by us to the speciali st, M. Guènèe, for his Speci es Gènèral . One will have thus all the Lepidoptera found to date in California. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 6 Now that M . Lorquin h as returne d to France we shall receive no thing more from this country; after him the spring will be dried up for a long time. One cannot count on gold seekers to collect insects." Boisduval goes on to descri be Lorquin's trip to the Philippines in 1857, via Hong Kong and Saigon and his subsequent return to China where he made a fine collection of Lepidoptera. He then return ed to Califo rnia bri efly, but w as unable to re sist the urg e to visit China and the Celebes again, he returned for nearly a year of exploration and collecting. Unfortunately, he followed Alfred Russell Wal lace by several y ears, and Wallace h ad already colle cted most of th e extraordinar y species of this South A sian area and had them described in England. One mention of the presence of Pierre Lorqui n at a regular meeting on April 1, 1867, of the Californi a Academy of Natural S ciences at San F rancisco is made in the Mining a nd Scien tific Press newspaper, published in San Francisco on A pril 6, 1867. The Minutes of that Academy meeting state that "Mr. Lorquin pre sented a case of but terflies from th e Philippines, Molucca, and Arrow Islan ds and Braz il." Lorquin had become a close friend of Herman H. Behr after meeting him in 1852 at the California Acad emy of Sciences. H is specimens were split a mong Boisduval (w ho received the great bulk of them), H. H. Behr, A. Guenée, A. R. Grote, and C. T. Robinson. Some specimens also went to the F elders in Vienna. Boisduval alw ays sent a specimen of each sp ecies when he described it from France back to Lorquin, and thes e co - types were turned over by Lorquin to Behr and preserved in the Museum o f the California Academy of Scien ces until the 190 6 earthquake and fire. Interestingly, Lorquin never learned English well enough to converse fluently in it (Grinnell, 1904). Befor e Lorquin arrived in California, very few butterflies had be en collected there. Of the la rge number of species of California Lepi doptera described by Boisduval (1852, 1859), all except five or six ( those being supplied by E. Doubleday) were collected by Lorquin. Boisduval lists 95 species of Lepidoptera that Lorquin took in Califo rnia, which included 83 butterflies. Some 53 butterflies were described as new species. One of them, Lorquin's Admiral, Limenitis lo rquini (Boisduva l), commemo rates the n ame of this ea rly collecto r. Especial ly appropriate is that this beautiful species occurs throughout much of the state. I t is interesting to note that Lorquin did not overlook c ollecting many of the really abundant and wide spread species of the 1850's. These included both new species and species already named by other workers such as Linnaeus, Hubner, and Lucas. The Lorquin Natural History Club, founded in 1913 and expired in 1919, and the Lorquin Entomological Society, founded i n 1917 and thriving to the prese nt date, preserve th e name of th is outstand ing early C alifornia lepidop terist. Jean Alphonse Boisduval (1799-1879) One of the mo st famous F rench entomo logists a nd a great syste maticis t in Lepidop tera, Boisduval was born in Normandy, France, on June 17, 1799, and died there on December 30, 1879. He produced some 50 publications, mos tly dealing with Lepidoptera. In Cal iforni a lepidopterology, he is most famous for describing and publishing the new Lepidoptera species taken by P. J. M. Lorquin in California, though he also described a number of other important insects na tive to North A merica and v arious Homopte ra and man y C oleoptera. Boisduval's gre at Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 13 Virginia and West Virg inia, a b uilder of ra ilroad s, a develop er of coal m ines, and ac tive in the community affairs throughout his life, he always found time to devote to the study of butterflies, and even more remarkably, to the preparation for publication of the results of his investigations. Edwards' first contribution to the pages of the Canadian Ento mologist appeared in the third number of the first volume in October 1868, and his 170th contribution to th at journal alone appeared in the thirtieth volume in January 1898. He also published many papers describing new butterfl ies in the Proceedings and Transactions of the American Entomological Society and the new, but short-lived, journal Papilio , edited by Henry Edwards. These two men were friends and corresponded with each other, Henry Edw a rds residing mostly in Californ ia during this time. There has be en some con fusion in th e litera ture because o f the simi larity in names. W. H. Edwards published his first book in 1847, entitled VOYAGE UP THE AMAZON , which gave an account of a tr ip that he made up that great r iver in 1846, not long after he gradu ated from college. His magnum opus, however, was THE BUTTERFLIES OF NORTH AMERICA. The first part of this great work was publ ished in April 1868 and the exquisite bea uty and detail of the plates a ttracted worldw ide attentio n. In July 1872, the f irst series o f parts forming a large quarto volume with 50 plates was completed. The second series of parts, containing 51 plates, was begun in May 1874, but not finished until N ovember 1884. The first part of the third series was issued in December 1886 and the last (the e ighteenth) in 1897, with some 51 plates. The quality o f the artwork an d the writ ing increa sed with e very part co mpleted . For example, in the th ird volume, n early half of the 51 plates are devoted to t he alpine or subarct ic species of the Satyrinae, and every species of North American Oeneis except O. taygete from Labrador, was figured wi th all the li fe stages fu lly describ ed. A w ealth of detai l on larval cha racte ristics wa s included, along with the eggs and pupae. Until Edwards began this mammoth work, little or nothing was known of the egg, l arvae, or pupae of any except a f ew of the commonest butterflies in North A merica. H e made the notable d iscovery in 1870 that eggs could be satisfactorily obtained by combining the female butterfly of any speci es with the growing hostplant of its l arva. He at once began having correspondents from throughout the country send him eggs procured in the field, in West Virgin ia, he could co mplete the ir rearing on substitute f oodplants an d accurately record egg, larval and pupal characteristics. This work was aided by the general extension of railroads across the co ntinent and he was able to secure eggs with reas on able speed from widely distant localiti es, including western states and terri tories. Mr. Edwards employed a number o f artist as sistants, a nd one of th ese, Mrs. Mary Peart, not on ly drew with extreme accuracy nearly all the plates, but also helped to r ea r a large numb er of the cat erpillars. T he coloring of the plates was done by Mrs. Lydia Bowen. In the first v olume of his great work, Edwards said "It is a matter of reg ret that, in so me instances, I shall not be able to say anything of the larvae. Even among our old and common species, the larvae are but little more known than in the days of Abbot, 70 years ago." In the Preface to the second volume, dated 1884, Edwards said "All this ha s changed, and today it can be said that the preparatory stages of North American butterfli es as a whole are better know n than all those of Europe." This was made p ossible by the important fact that Edwards uncovered about the females, as related above. In his own words: "In 1870 I discovered an infallible way to obtain eggs from the female of any s pecies of butterfly, namely by confining her with the growing food- plant. If the eggs mature, they will be laid" (Anonymous, 1909) Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 14 At the age of 75, Edwards gave up his studies of butterflies, feeling apparently that his advanced age precluded him from working with the ability and success that he had s o remarkably displayed. However, he continued working in literary areas , on such s ubjects as Shakespea re and a geneology of the Edwards family; th e latter became his last work, published in 1903. During the period of his active studi es on butterflies, new sp ecimens of North Amer ican butterflies received by the Smithsonian Institution in Washington were sent to him for description and classification. Additiona lly, the Impe rial Russian Governme nt and the B ritish Museu m sent to him fo r examination and description many North American examples that appe ared to be new. As a result, he becam e the autho r of a large number of new species, inclu ding many whose ranges in western North America include California. The Carnegie Museum purchased the Edwards collections before his death and the specimens were curated by Willia m J. Holland, the direct or, who used them as the basis for his own book, THE BUTTERFLY BOOK (Bethune, 1909). While most o f the specime ns from the great E dwards collect ion went to the Carnegie M useum, some ended up in other institutions by exchange or trading for life history material (e.g., Irwin, 1966; Irwin, 1972). Edwards had extensive correspondence with many of the leading professional and amateur lepidopterists in the U nited States. Some of this extens ive correspondenc e has been analyzed by his biographer, F. Martin Brown (e.g., 1958, 1959). This being before the days of carbon paper or xerox machines, co pies of the letters were frequen tly made by pressing a moist pie ce of tissue paper again st the origin al letter, resulting often in s meared copies o r (if the tissue was too dry) only a bit of a let ter here and there making a bold tra nsfer. Nevertheless, the correspondence between Willia m Henry Edwards and such people as Spencer Fu llerton Baird offers c onsiderable help in fix ing type loc alities an d distrib utional r ecords from the early c ollectors in the wes tern s tates. In particular, F. M. Brown has published a monumental survey of all the Edwards types and used his indefatigable energy to trace down existing correspondence, hints on specimen labels, etc., to recr eate the often obscure story of w here these spe cimens were colle cted and how they reached Edwards for description. During his lifetime, Edwards wrote about 265 sci entific papers, mostly on butterflies, wherein he proposed so me 356 names for nea rctic butterflies as w ell as for some moths (dos Passos, 1951). Cyril F. dos Passos, himself a famous lepidopterist of the 20th century, had the good fortune to discover in 1949 an autobiographical m anuscript written by Edwards and completed three months before his 80th birthday. These notes were published by dos Passos (1951) and a reading of them reveals many o f Edwards' earl y experi ments with life his tory rearings, illustrations, and of course his wealth of correspondence. At Edwards' urging, friends experimented with the effect of cold on pupae of Papilio , especially m arcellus , which showed the effects that cold and warmth cou ld have on coloration and pattern development. He also discovered between 1875 and 1878 that the larvae of the blue Celastrina pseudoargiolus are attended by ants, and he noted the intimate behavior of the ants in proximity w ith the last segments of th e larvae. Subsequen tly, he and a colleague discovered the sugar - secreting fluid glands on the back of a larva, which served as sig nals for the ants to come to segmen t eleven. Edwards then sought to find out why the ant s would be of any use to the l arvae, and the follow ing summer h e watched the larvae (tended by ants) in the fi eld. He discovered that the ants protected the larva from ichneumonid w asps. These and many other discove ries were made as a result of his exhaustive life history work. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 15 In terms of California species, we know from Edwards' correspondence and publications that in April 1886, he received eggs of Apodemia mormo from William G reenwood Wright at San Bernardino, Califo rnia, and he received eggs of many species from C. F. McG lashan at Truckee, California. In 1889 and 1890, he r eceived eggs of many southern Califo rnia species from Wright at San Bernardino. Edwards noted in his autobiography that he became a correspondent with the actor Henry Edwards about 1872 and from that time until Henry Edwards' death, they frequently exchanged letters and butterflies. They first me t personally in December 1887 at Henry Edwards' home in New York where they spent the morning in his study. What an inspiration it would be to read a transc ript o f their conve rsation s! Theodore L . Mead ( 1852-1936) Theodore Mead was born in the Cats kill Mountains of New York on February 23, 1852. His forebea rs had imm igrated from England over 200 years earlier. His early yea rs were fil led with adventures in natural history and when butterflies took his interest, he becam e acquainted with William H. Edwards. Later, he was to become W. H. Edwards' son- in- law. In 1866, he was elected to the American Entomological Society and he spent the following summer i n West Virginia working with Edwards. Apparently, the young man's enthusiasm and ability s truck a responsive chord, for in 1871 Edwards arranged that Mead accompany the Wheeler expedition to Colorado as a paying member. Edwards shared Mead's expenses and also received a portion of the catch from Mead (Mead, 1875). A fter he collected in Colorado, Mead continued westw ard in the fall of 1871 and visited entomologists in California. He the n spent some years at Cornell University earning a degree in civil engineering. In 1877 he sold his collection of b utterflies to the Carnegie Museum at Pittsburgh, and from that time on he devoted more of his life to botanical and horticultural work. However, in 1878 he di d take a second entomologically- oriented trip to Californi a, arriv ing in S an Franc isco in ea rly May and leaving there on May 7 for Los Angeles and the San Gabriel V alley. On this California trip, Mead v isited the Mojave Desert, Yosemit e Valley, Big Trees, Mariposa County (Essig, 1931), Summ it Station on Donner Pass, Lake Tahoe, and Freel's and Talloc peaks, the la st area be ing where he collected the types o f Oeneis ivallda Mead. On h is way home, he collected at Virginia City, N evada, and at several poin ts in Utah and Wyoming. In 1881, Mead joined Henry Edwards and A. R. Grot e in starting the journal Papilio , which was published for four years. In 1882, Mead married the daughter of William H. Edwards.and after a honeymoon in England, he s ettled in central Florida to try his han d at orange- growing. From this time on, he pursued botanical interests exclusively and did not publish further on butterflies. He died in Winter Park, Florida i n 1936, shortly after publishing his autobiography (New, 1935; Brown, 1955). While most of M ead's lepidopterological fame (Brown, 1934) is from his collecting in Colorado, a period which supplied many specimens to W. H. Edwards for description, he did collect significantly in California and helpe d launch many papers on the western butterfly fauna in his joint founding of the journal Papilio . Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 16 Richard H. Stretch (1837-?) A pioneer lepidopterist on the Pacific Coast, and an associate of H. H. Behr, Henry Edwards, Samual H. Scudder, and William H. Edwards, Richard Harper Stretch was born November 25, 1837 in England. He developed an intense interest in natural history very early. Following his boarding school years, he became an apprentice in a dry goods shop. In 1861, he received an invitation to visit relatives in Illinois, but returned to England after about a month. In 1862, he visited his uncle once again in Illinois, making extensive collections there, and then in 1863, he joined an emigrant party bound for California. His small band of five persons was able to pass unmolested through hostile Indian country despite the fact that other wagon trains were attacked and wiped out at the same time. Their protection apparently ensued from Stretch's entomological pursuits. When Stretch displayed his preserved insects to a friendly Indian at the outset of the journey, the word was passed among the Indian bands that a big me dicine chief was coming and that no harm should befall him. When he arrived in Utah, Stretch changed his plans and went south into Nevada. He proceeded to begin a career in Virginia City, then a rip-r oaring frontier mining camp. First employed as a cook, he later ran a sawmill and after that became associated wi th a partner in a land office business. His entomological knowledge served him well, for he was able to discriminate minute differences and grade with ease the various types of ore. To his astonishment, he was elected State Mineralogist of Nevada. Soon he became known as the best maker of mining maps in America. In 1867, Stretch journeyed on to California. En route, he was among the earliest of the emigrants to cross the fearful Death Valley region. He visited San Francisco and also descended into Chihuahua, Mexico. In the winter of 1868-1869, he returned to Ne vada to Virginia City for the U.S. Geological Survey to make a study of the famous Comstock Lode. He returned to San Francisco and became city surveyor in 1870. Despite all the pressures of business on these trips, his interest in entomology never flagged and he discovered many new and interesting species in both California and Nevada. His most significant California collecting began in 1875 and continued through 1876, when he lived and collected extensively in the Havilah area in Kern County. Here he discovered the dainty blue Philotes speciosa , which was named by Henry Edwards. Henry Edwards also described a number of other butterflies taken by Stretch at Havilah. In 1888, he moved to Seattle, Washington, and from there, various mining jobs took him to nearly every state in the West, as well as British Columbia. Shortly after his wife died in 1885, Stretch donated his entomological library to the Mechanic's Institute in San Francisco and his magnificent collection, replete with many types and rarities and numbering about 13,000 specimens, was given to the University of California at Berkeley (Coolidge and Newcomb, 1920). Essig (1939) later estimated Stretch's collection at Berkeley as numbering about 5,000 specimens. Stretch is probably most remembered in entomology for his personal work on the moths, including his classic work "Illustrations of the Zygaenidae and Bombycidae of North America," published in 1872-73. While he knew many of the other great early entomolog ists of the West and people such as Thomas Edison and Mark Twain, Henry Edwards was Stretch's closest friend and companion, and his entomological studies practically ceased when Henry Edwards died in 1891. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 17 Oscar Theodor Baron (1847-1926) Baron was born on September 9, 1847, in Germany, the son of a schoolmaster. He studied collecting insects at the early age of 13 years. As a young man, he was shipped as a sailor to the Orient and visited India, China, and the Austra-Asian region. From Java, he shipped to the west coast of America where his first job in the New World was as a surveyor. The little we know about Baron's life after that has been augmented by the researches of F. M. Brown who examined the extensive correspondence that Baro n had with Henry Edwards and Herman Strecker. Brown (1965) was able to reconstruct a year-by-y ear itinerary of Baron from the time he was 28 years old (1876) and working at loading schooners applying between a small mill in Mendocino County and San Francisco. In his spare time Baron was collecting mo ths, butterflies, and other insects. He knew a fair amount about saturnids and other moths in 1876 and 1877, and apparently began exchanging butterflies with W. H. Edwards at this time (although no account in Edwards' journal survives). He also exchanged specimens with Henry Edwards and H. Neumoegen. In 1879 Baron set out from Navarro in Mendocino County on a long collecting trip, starting in mid -April. By September, he returned with a good catch from the summer season. In subsequent years, he traveled as far as San Diego in southern California, but most of his California collecting was in the north -central counties. In 1882, he met William Greenwood Wright and instructed him in how to collect and preserve Lepidoptera. Letters in the Wright collectio n and in the Pacific Coast Entomological Society files show that Wright started on his long career as a lepidopterist at this time. In 1884, Baron apparently traveled to Mexico and then later to Ecuador. By 1887, he was back in southern California and also working with a crew of engineers in Pima County, Arizona. He collected some butterflies and butterfly eggs for W. H. Edwards during this time. By 1892, Strecker inquired about purchasing Baron's collection, Baron estimated that there were perhaps 27,000 Lepidoptera, of which 6,000 were from California. Baron did not reach an agreement with Strecker, and sometime in 1893, he returned to Germany, and lived there for the last 30 years of his life. He sold a large part of his collection in England en rout e to Germany but retained about 100 drawers of American butterflies for his own use. Some of the specimens that Baron shipped to Strecker are in the Chicago Natural History Museum. Those that remain from Neumoegen's collection are in the United States National Museum, and Henry Edwards' specimens from Baron are in the American Museum of Natural History. Many of the specimens in the W. H. Edwards collection at the Carnegie Museum bear on their labels "O. T. B." in Edwards' handwriting, testifying to their origin (Brown, 1965). James Behrens (1824-1898) James Behrens was one of the pioneer collectors in California. Behrens was born in Germany on June 30, 1824, and died at San Jose, California on March 6, 1898, at the age of 74 years. He was considered one of the best early collectors in California, acqui ring specimens of all orders but describing few species himself. The Lepidoptera he collected were described by many moth specialists as well as W. H. Edwards. His large personal collection was given to the Museum of Lubeck, Germany, a few years before his death (Essig, 1972). Among the California Lepidoptera named for him is Argynnis behrensi Edwards, now called Speyeria zerene behrensi (W. H. Edwards). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 18 Herbert Know les Morrison (1854 -1885) M orrison was a notable insect collect or who was born at Boston, Massachusetts on Jan uary 24, 1854, and died at a very young age on June 15, 1885. He publis hed principally during the years 1873 through 1875, and chiefly in the Proceedings of the Boston Soci ety of Natural History . He made a number of impo rtant trips through the western states as well as in the sou theastern U.S. In 1874, he collected insects in Colorado, in 1878 in Nevada, Utah and the Black Hil ls of what was to be South Dakota, in 1879 in Washington territory, and in 1880 in Arizona and southe rn California in 1882. He re turned to Arizona and New Mexico in 1883 and the n went back to the Southeast where he died in the spring of 1885 at Key West, Florida, from an attack of dysentery. He was s aid to be an exceptionally capable professional collector, with great physical strength and stamina. Mann (1885) comments that Morrison would frequently walk forty miles a day in pursuit of insects, and then would take care of them that evening while capt uring moths th at were attra cted to his light. He supplied specimens of principally Lepidoptera for systematicists in both A merica and Europe. He also per sonally named a number of species from the western region. Species of Lepidoptera were named for him by Henry Edwards, W. H. Edwards, H. Druce, A. R. Grote, and C. V. R iley. Amon g the C alifornia butterfl ies, Anthocharis cethu ra morrisonni was named afte r him by William Henry Edwards, and may st ill be found today in the San Joaquin Valley (Anonymous, 1885; Essig, 1972; Mann, 1885). Edward Palmer (1831-?) Dr. Edward Palmer was born in England on J anuary 12, 1831 (though at vari ous times he gave the date of his birth as late as 1837!), and he arrived in the U nited States in 1849. He studied homeopathic med icine and also s tarted a career a s a botanical colle ctor. W hen the Civ il War bro ke out, he joined the army in Denver, Col orado, and served as a surgeon in the ar my at various posts in the Southwest. In 1865, he was posted to Fort Whipple in Arizona and began collecting insects for the Smit hsonian Ins titu tion and for private collectors. He left the army sometime around 1868 and beca me a profes sional f ield nat uralist, o ften employ ed by the D epartmen t of Agricu lture to explore for plants in the Southwest and in Mexico (Brown, 1967). The types of m etalmark butterfly, Apodemia palmerii (Edwards), were collected on a trip in 1870 through Utah and Nevada, including a visit to the mouth of the Colorado River south of the boundary of California and Arizona in Mexico. In the mid - 1870's, he collected in southern California, including the San Diego area. He made a maj or trip in 1877 into southern Utah and adjacent nort hwestern Arizona, collecting insects in earn est for the m useum at H arvard Un iversity. The Lepid optera co llected on that trip came to Samuel H . Scudder at Harvard, including four new species: Neominois dionysus , Anthocharis thoosa , and Pholisora libya , a ll described by Scudder in 1878, and an unnamed species of Erynnis . Whi le no species col lected by Dr. Palm er in southern California were named as new species, the ones he co llected outside of our state such as Apodemia palmeri and Anthocharis sara thoosa and Pholisora libya reach our territory. In fact, the typ e locality of Pholisora libya and possibly of Apodemia palmeri is thought to be Littlefield (the modern name of Beaver Dam, Palmer's locality name), Mohave County, Arizona, about 38 miles s outhwest of St. George via U.S. Highway 91 in desert country (Brown, 1967). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 19 Samuel Hubbard Scud der (1837 - 1911) Born in Boston, Massachusetts on April 13, 1837, Scudder has been regarded as one of the greatest systematic ent omologists in Ame rican biology . Unti l he died at Camb ridge, Massachusetts on May 17, 1911, he pursued three lines of fundamental work: insect paleontology, systematics of Orthoptera, and systematics and morphology of Lepidopter a. He attended both Williams College and Harvard for his B.S. degree (received in 1866) and had al ready become an assistant to the g reat zoologist Louis Agassiz at Harva rd some two years prior to his degree, in 1864. In 1879 he became Assistant Librarian of Harvard University and continued in that post until 1882. He was the founder and early editor of the j ournal Psyche , and P resident of the Boston Society of Natural History from 1882 to 1887. In 1886 t hrough 1892, he was paleont ologist for the U. S. Geological Survey and in that position laid the foundation for the study of fossil insects and became the greatest Americ an authority on insect paleontology (Essig, 1972). H e described 838 Terti ary insect s from the fo ssil deposits o f North A merica, la rgely from F lorissan t in centra l Colorado. He also named 80 species of b eetles from Pl eistocene deposits o f Canada and Massachusetts as well as nearly 20 0 species of Pa leozoi c insects. In 1871, he published the Index to the Known Fossil Insects of the World in a 744 - page issue of the U.S. Geological Survey Bulletin . His reputation for his work on Lepidoptera began in 1889 when he published the great three- volum e work entitled The Butterflies of the Eastern United States and followed t his treatise with several minor papers on butterflies over the next four years. In all, he published 791 scientific papers while pursuing his three distinct lines of work simultaneously. His output was so prodigious and influential on the growth of American entomology that, although he never personally visit ed the West, his name is as f a miliar to eve ry entomolog ist in the W est as it is throughout the entire country (Essig, 1972). He named a number of western butterflies, including Pieris napi marginalis (Scudder), Anthocharis sara thoosa (Scudder), Hesperia juba ( Scudder), Erynnis ic elus (Scudder and Burgess), E.propertius ( Scudder and Burgess), E. f uneralis (Scudder and Burgess), Thorybes mexicana nevada (Scudder), and Hesperopsi s libya (Scu dder), which occur in Ca lifornia . Tyrone Reakirt (1844- ?) Tyrone Reakirt was very much a mystery person t o most biographically inclined entomologists. The short sketch of his life by Essig (1972) is declared to be complete erroneous by Brown (1964), who made an exhaustive search for letters from Reakirt in order to learn more about the man. Reakirt was born in Philadelphia, Pennsylvania on April 21, 1844. He went int o partnership eventually with his fat her and brother Daniel in the i mporting and wholesale drug business. He star ted a new venture i n Delaware in 1868, w here he manufactured w h ite lead and, later, lead acetate. He became a member of the E ntomological Society of Philadelphia in 1863, at which time he w as 19 years old, and he h ad apparently some financ ial substance at that time. He Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 20 subsequently became well - known as a student of the butterflies of the American tropics and of the Indo- Austral ian region. He also pub lished th e first su mmary of o ur knowle dge of th e butterflie s of the Rocky Mountain region in 1866, based on the c ollections brought back from Colorado by James Ridings in 1 864 as well as tw o earlier coll ections made by Willi am Wood and Winslow J . Howard th at were in the Socie ty of Phila delphia' s collecti on. Inte restingly, W. H. E dwards considered Reakirt to be the North American Rhopalocerist most competent to h andle the very large colle ction of bu tterflie s sent to the Smi thsonian Institut ion from Co lombia (Bro wn, 196 4). Reakirt's co llection ul timately was combined with Streck er's and was d eposited in th e Field Museum in Chi ca go. Reakirt apparently did very little, if any, field collecting and instead purchased his specimens from Lorquin and others. In November 1868, for example, he wrote to Strecker and told him that he was expecting a shipment of 1,000 specimens of butterflies from the Philipp ines, Molucca and California. During the period of 1866 - 1868, Reakirt employed Strecker to mount much of the material that he was receiving, and in the summer of 1868 he turned his collection over to S trecker in orde r to sell it to rai se a dditional funds. The original price was set at $1,800 and Strecke r held the co llect ion for some mon ths und er agreement to s ell it fo r Reakirt. R eakirt eventually offered the collection to Strecker but, though Strecker agreed to purchase the coll ection fo r $1,446, he made none of the st imulated payments. Reakirt got into serious fi nancial trouble and his company was forced into bankruptcy, so he fled the country early in 1871 to Lima, P eru. Strecker received 1 1 lett ers from th e exile d uring th e next ye ar and then n othing further is known about th e rest of his life . His name is co mmemorated in the butterflie s he nam ed and tha t were named after him such as Pa pilio indr a indra Reakirt, Pontia occidentalis (Reak irt), Lycaena mariposa (R eakirt), Leptotes marina (Reakirt), Oarisma g arita , and a patronym, Anthocharis sara form "reak irti," the ea rly spring form of ou r common Cali fornian o range -tip. James John Rivers (1824-1913) Rivers was born in England on January 6, 1824, and died in Santa Monica, California on December 16, 1913. During his nearly 90 years of l ife, he personally knew Charles Darwin, Alfred R. Wallace, Thomas H. Huxley, and other naturalists then active in the Entomo logical S ociety of London during their college days. In about 1867, he came to the United States, moving from Kansas to Colorado to finally California sometime between 1875 and 1880. He joined t he early group of naturalists asso ciated with the Cali fornia A cademy of Scienc es, including H. H. Behr and James Behrens, and ser ved as curator of Organic Natural H istory in the University of C alifornia until 1895, when he moved to southern California to live until his death. While he collected very widely in diverse groups of animals, he was inter ested in Lepidoptera, particularly the genera Euphydryas and Melitae a (Essig, 1972). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 21 In California and the ot her Pacific states, John Adams Comstock was the most fa mous lepidop terist of the first half of the twentie th centu ry. His life h as been re counted by many au thors (e.g., Emmel and Emmel, 1973, 198 9; Martin, 1971, 1972; Van Doren, 1983), from which the following highlights are taken. During his 87 ye ars, he published more than 200 scientific papers on the life history and taxonomy of wester n Lepidoptera and published the outstanding seminal work on C alifornia butte rflies, Bu tterflie s of California , in 1927. His ea rly field work throughout California and his extensive correspondence and cooperative endeavors with othe r outstanding lepidopterists in the United States and around the world richly earned him the title of "Dean" of the Califo rnia lepid o pterists. His 1927 book has stood for more than half a century as the preeminent authority on the biology and ecology of the California butterfly fauna. It is only in t he 1980's that several new works have begun to appear and to amplify th e outstanding foundations laid by Dr. Comstock in his pioneering book. It is appropriate to open this account of the life of a great lepidopterist by going back 68 years to Comstock's (1920) preliminary published announcement of his project to prepare a book on the bu tterflie s of Cali fornia, wi th the c olor illu strations to b e executed in the ou tstanding "three- color copper - plate process." The plates for this work were prepared in Los Angeles under the direction of Mr. Raymond Thorpe and at the time they represented "the highest expression of this form of reproductive art. T hroug h their u se it wi ll be po ssible to identify any specime n captu red in the state at a glanc e." Comstock h ad this announcement an d these color plates p ublished with related papers on various butterfly genera in a dvance of the book in the quarterly issue s of the Bulletin of the Southern California Academy of Sciences over the subsequent half -dozen years, in order to "stimulate the study of this most fascinating branch of the natural sciences, and bring new recruits into the Academy's 'Outdoor Army.'" But his closing paragraph in his 1920 announcement conveys perhaps better than any other words why his outstanding 1927 book made the marvellous Californ ia butterfly fauna ha ve such last ing inter est to lepido pterists : "Califor nia is unusu ally favor ed as an en vironmen t for this study. With in its co nfines are found nearly t wo hundred and f ifty disti nct species of butte rflies, not count ing the many interesting variet ies, local races, and aberrations. The d iversity o f our cultivated flora gives a footing for many introduced species. Our isolated mountain ranges have developed extremely local for ms. In a word, California has compressed in its con fines the elements of an entomological empire -- a veritable 'happy hunt ing grou nd' for th e butterfly enthu siast." The Comsto ck family t raced the ir ancestry b ack to Wil liam C omstock, w ho emigrat ed to Massachusetts from England in 1635. John Adams Comstock himself was born on January 30, 1883, in Evanston, Illinois. He was one of seven childre n in a wealthy family. His father , John Adams Comstock, Sr., had been born O ctober 2, 1845, in Franklin, Oakland County, Michigan. His mother was Cornelia Hurd who was the daughter of Judge Harvey B. Hurd, one of the founders of Northw estern Unive rsity and an eminen t scholar an d ju rist. This maternal grandfather , Judge Hurd, lived with the famil y for some years and contributed toward making the Comstock hom e an intellectual and cultural center which resulted in even greater benefit to the young Comstoc k than his early formal ed u cation. Young Comstock was fully trained in the fine arts of natural sciences by individual tutors coming to his home during high school and coll ege. He also took classes at the Chicago Art V. THE GOLDEN E RA OF CALI FORNIA LEPIDOPTERA DISCOVERY (1890 -1 940) John Adams Comstock (1883-1970) Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 22 Institute and the Laflin and F ield Museums of C hicago. He o nce said of t hese early influenc es, "It was the wi sh of ou r family that I s hould fo llow art a s a voca tion, but my tastes were more inc lined toward natural history. My insect collection was begun at the age of twelve." In 1901, John Comstock and his brother Hurd t raveled to Lake Josephine and Avon Park, in south central Flo rida, to collect bu tterflies . Some of these specimens a re still in the collection of the Natural History Museum of Los Angeles County. T he following year, in 1902, he and his brother and a Professor Snyder went to Hall Valley in Colorado to collect mountain species. This locality had first been made famous by David Bruce's collecting for Henry E dwards, some thirty years earlier. By the age of 15, John Comstock was Secretary of the Chicago Entomological Society and became its Recorder at age 19. In 1904, at the age of 21, he became an architectural designer for Myron Hunt in Chicago. During the summers he taught nature study in Lake Geneva, Wisconsin. In the fall of 1905, Comstock m oved to East Aurora, New York, with the intention of improving his natural ability as a designer and w riter. For the next several years, he j oined the Roycroft Arts and Crafts Community. This comm une had been founded just prior to the turn of the century by Elbert Hubbard, and the community was based on the teachings and philosophy of John Ruskin and William Morris in England. During this time , he conducted nature study classes and introduced nature - inspired designs into the Roycroft work. He improved on his fu rniture and bookplate designing as well as bookbinding and illustrations, and also delved into metalwork and jewelry design along with other creative work. The Roycroft Press was a key part o f this community and Comstock learned much about book design, bookbinding, il lustration, and bookplate design at that time. In his subsequent two books on butterflies and t he Comstock geneology, the influence of this experience in New York is q uite appare nt. He marri ed Carolyn Townsend in 1905 and they had three children: John Sterling, Jean, and Bette. The family moved to Santa Rosa, California, late in 1906, and by 1908 he and his sister had esta blished the Craft- Camarata ar ts and crafts shop. H e als o worked for Luther Burbank as a designer for Burbank's nursery publications. By 1910, he opened a nother Craft - Camarata shop at Santa Barbara , Californ ia. In the su mmer of 1912 , he taught at the Carme l Summer S chool of A rts and C rafts. In the fall of 1911, he s tarted medical college in Los Angeles, enrolli ng in a four - year course at the College of Osteopathic Physici ans and Surgeons in Los Angeles. In 1915 he passed the State Bo ards and was licen sed to practice medicine and su rgery in Cal ifornia. Here h e met and married his second wife, Carolyn, who was also a doctor. Later, Com stock's D.O. degree was changed to an M.D. This license was kept up during the rest of his life, and he pr acticed medicine to earn su fficient income for th e pursuit of h is ot her interes ts. His interest in natural science and especia lly entomology nev er waivered during these years of medical study. He collected w henever time permitted, and especially continued his interest and studies of West Coast Le pidoptera. In the early years of the So uthwest Museum, whic h was placed on a hill overlooking the current Pas adena Freeway in the Highland Park portion of Los A ngeles, he became Honorary Curator of Entomology. In 1920, h e became the Southwest Museum's Associate D irector and then its Dire ctor f rom 1921 to 1926. Comstock simultaneously helped to build the Southwest Mu seum during years of severe financial trouble, enhance i ts collections, and establish firm public support for t he Museum. In 1926, the natural history sections of the Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 29 discovered, etc. Skinner apparently sent him many references and data on various names (Gordon F. Pratt, i n litt.). Jean Daniel G under (1888 -1948) J. D. Gunder was born in New York C ity in 1888, but moved to Pasadena, California, over 25 years prior to his death on Nove mber 17, 1948. He was keenly inte rested in butterflies of western No rth Americ a, part icularly Californ ia, and he descri bed some 21 2 species, races, forms, and aberrati ons in a series of papers in the leading entomological journ als of America. H e is most well - known for his outstanding systematic work on checkerspot butterflies, as published in his paper "The Genus Euphydryas of Boreal America" in 1929. With ext raordinary insight and taxonomic acuity, he managed to reduce the rampant systematic confusion in that genus, with its tremendous geographic variation, into just seven specie s. He used genitalic structures in this paper to such an extent that it has become one of the landmark papers that has shaped our modern classification and concept of butterfly species, as L. P. Gray (1970) has stated. Gray (1970) also emphasizes that Gunder was doing equall y brilliant work with the genus Argynnis (now called Speyeria ), although this work was not finished before his active interest in butterflies was dropped in 1935, when economic conditions forced him to give up his collection and enter a new business (Martin, 1948). In 1937 Gunder sold his collection of about 28,000 specimens to the American Museum of Natural History in New York, while his outs tanding entomological library was acquired by Cyril F. dos Passos. In the latter years of his w ork, Gunder concentrated on h is princ ipal in terest o f aberrations and "transition forms," a fascina tion which unfortunately led to his disparagement by certain eastern lepidopterists who dismissed Gunder as a wild amateur (e.g., Gunder, 1927). It is particula rly unfortuna te that his pioneering systematic r evisions of Euphydryas and his incipient revision of Speyeria were overlooke d for some time as a result of that attitude. During Gunder's career, at least six butterflies and two moths were named in his honor by leading entomologists. John F. Garth (1909- ) For more than 45 years, John S. Garth has been a princi pal researcher and important contributor on the butterfly fauna of western North America. His books, monographs and scientif ic papers on th e butterfli es of Califo rnia and of our major w estern nationa l parks are familiar to all America n lepidopte rists. In l98 6, he co- authored a major new work entitled Californ ia Butterflies , pu blished by th e University of Californ ia Press, w hich stimulated addition al wide public interest in our butterfly fauna and which encouraged many new students to take up the study of California butterflies. For these and m any other accomplishments in lepidopterology, Garth was honored by the Pacific Coast Section o f the Lepidopterists Society with the John Adams Comstock Award in 1987. John Shrad er Garth is that r arity of citizen: a native Calif ornian. He was born in Los Angeles, California, on October 3, 1909, and grad uated from Long Beach Polytechnic High School in l927. He began his first butterfly collecti on at the age of nine, collecting in Bixby Park. His Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 30 early in terest in souther n Californ ia Lepid optera w as foste red by cont act at th e Los An geles County Museum of Natural History w ith John Adams Comstock and Lloyd M. Martin, w ho soon became life - long friends. He was also to meet many others of the early sou thern Califo rnian lepidopterists through the monthly meetings and field trips of the very active Lorquin Entomological Society. Following high school, John Garth entered the University of Southern Ca lifornia where he was to take his next three degrees: a Bachelor of Music degree in l932, a M.S. degree in l935, and a Ph.D. in Zoology in l941. He also married I sla L. Detter on June 25, l940, to whom he is still happily married and who is a frequent participant with John at the Paci fic Slope Section meetings each year. Mixed in these acad emic years of study at U.S.C. were summer sessions at the Yosemite Field School (1933), Cornell University (1937), and the University of Pennsylvania (l940). The multifaceted foundations to Garth's lifelong series of accomplishments and expertise in a diversity of fields -- from music to entomology to crab systematics and biogeography -- were laid during these key years. In fact , it was Garth's outstanding ability in music t hat gave him his first gre at opportuni ty for travel and f ield resear ch in the tropic s. In l93l, when Garth was a senior at U.S.C. and an instructor at the Long Beach Institute of Fine Arts, he was recommended by the Dean of the U.S.C. College of Music to go w ith Captain Allan Hancock of Los Angeles on a scientific expedition to the eastern Pacific. Garth was studying piano, and his interest in butterflies was enough to satisfy Hancock who was looking for someone with a background in the natural sci ences to play in his shipboard ense mble during the trip. By just such serendipity, events dr ew John Garth to his life's work and f orged his lifelong associ ation with Captain Hancock, a great philanthropist, musician, real estate developer, mariner, aviator, scientis t, industri alist, far mer and fou nder of wh at became th e United C alifornia B ank. Thus in the middle of the Great Depres sion, Garth left school at the age of 2l to embark on this thr ee -month- long, l2,000 - mile voyage with Hancock and his group of research scientists from major unive rsities. The boat was the V elero III , a l95 - foot motor cruiser which H ancock had outfitted for the first scien tific expedition und er his auspi ces. They sai led for the Galap agos Islands off the co ast of w estern South Ame rica to stu dy marine life and ocean cur rents, w ith some terrestria l fiel d work on the insects as John's planned assignment. Hancock played the cello, and as part of this patron's expedit ion program, the musical ensembl e played classical music ea ch evening after the day's work. This init ial voyage started the Hancock Pacific Expeditions, which were conducted annually from l93l to l94l aboard the Valero III . John's career as a marine biologis t started with th e first voya ge and he as sumed increa sing shipboa rd respons ibilitie s with each successive vo yage. In l940, the Allan Hancock Foundation was establi shed on the University of Southern Californ ia campus to serve as a reposito ry for mar ine colle ctions gathe red from M exico, Cent ral and South A merica, and the Galap agos Islands, an d Garth served successively a s Research Associate, Curator of Crustacea (i ncluding ot her arthrop ods), Ass ociate Profess or of Biolog y, Professo r of Biologic al Sciences, and Chief Curator of all the col lections u ntil his forma l retiremen t in l975 . He co ntinued as active ly as ev er the reafter as Professor Emeritus at U.S.C. He published more than 50 scie ntific papers and monographs in entomology and crustacean biology, and is widely recognized as a pre - eminent world authority on cra b taxonomy, distribution and Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 31 ecology. In the course o f his academic career, he guided more than 40 g raduate students towards advanced degrees, and is known as the key scientist who produced a whole generation of Ph.D..- level carcinologists. Through the Hancock Pacific Expeditions, John Gart h was exposed very early to the riches of the t ropics and their b ewildering d iversity of Lepid optera a nd other o rganisms. He part icipated in expeditions to the Galapagos Islands in l93l - 32, l933, l934, l935, and l938. He visited mainland Ecuador in l934, Peru in l935 and l93 8, and the Caribbean islands of Aruba and Curacao (Dutch), Trinidad and Tobago (then British), and Colombia and Venezuela in l939. Additionally, he visited Mexico, the Marsha ll Islands , India, Ceylon, and the Maldive Islands from the l940's to the l960's. He also travelled to many European m useums, Australia, New Zealand, Singapore, Mala ysia, Thailand, Hong Kong, and Japan during the course of his scientific researches. Wi th all this field work in exot ic places, Garth's first love of Lep idoptera was alw ays centered in th e western United States. B ut it w as not ea sy to im mediately start pr ofessional work on w estern L epidopte ra in tho se days, and though John publi shed several important papers in the early l 930's on Yosemite National Park butterflies and the geographical race concept in Lepidoptera, he looked for additional entomological background in his usual thorough manner. While John Garth majored in zoology at U.S.C ., with a strong emphasis on marine invertebrates, his early interest in entomology ha d to be carried on as a minor. Unable to obtain graduate- level work in entomology at U.S.C. in the l930's (entomology was re presented on his Ph.D. committee at U.S.C. by John Adams Comstock from the Los Angeles County Museum across the street), G arth sought s ummer cou rse work in th e East, fi rst at Corn ell (l937 ) and later a t Pennsylvania (l940). He also met E. O. Essig, a famo us wes tern en tomologist, as a me mber of the Yosemite School of Field Natural History in l933. During t he Second World War, John was assigned to t he Army Air Corps, w here he serve d as lieutena nt/captain at Boca R aton Army Air Base in Florida, with responsi bility for mosquito control. A ccess to a military reconnaissanc e car allowed him to collect Lepidoptera extensi vely in Broward and Dade counties, although gasoline rationing permitted on ly two priv ate collectio n trips (to Ma rco Island on the Florida west coast, and to Marathon in the Florida Keys). He also renewed his acquaintance with J. H arold Mattheson in Miami, who showed John many of his choice collecting localities. Earlier during the Hancock cruises, Garth had occasion to visit B arro Colorado Island in Gatun Lake, Canal Zone, where h e met many i nfluentia l scientists of note fro m the American Museum of Natural His tory such as Frank M. Chapman (ornithologist and author of well - known books on the tropics), Frank E. Lutz (eastern entomologist and autho r of The Insect Boo k ), James S chwartz (hy menopteris t), and Irv ing Huntington (lepidopterist). Thus, while Garth was "on his own" as entomologist with th e Hancock cruises, and at that time had had no formal work in entomology, these contacts with professional entomolo gists at B arro Co lorado (a nd with med ical ento mologists at Groga s Memorial H ospital in Balboa) proved helpful in answering questions that arose concerning tropical insects and entomolo gy in general. Some of his first rain - forest collecting was done along the trai ls of Barro Colorado Is land, an d it was here that Ga rth collected his first iridescen t blue Morpho amathonte . In the summ er of l933, Garth at tended the Y osemite S chool of F ield Nat ural History with l9 other p articipants, and his inst ructo rs included notable sc ientists fro m Stanford , Berkeley , and the National Park Service, including Dr. Harold C. Bryant, co -found er of the school and at that time assis tant directo r of the N ational Park Service. T he School included a high- country trip across Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 32 the Sierra Nevad a and the first scien tific survey o f a remote area along Yosemite Creek. G arth's lepidopterological discoveries during this summer were published over the next several years in the Bulletin of the Southern California Academy of Sciences. In the summer of l935, Garth was app ointed seasonal ranger - naturalis t in G lacier Nati onal Park, Montana. Stationed at the former St. Mary's Lodge on upper St. Mary's Lake, his duties included morning and afternoon nat ure walks, evening boat trips on the lake, and an eve ning nature talk each night at St. Mary's Lodge. This pace continued unabated seven days a week for six weeks straight be fore he was g iven a week off! He p romptly used th is time to drive to G reat Fa lls, where Dr. C. C. Albright showed Gar th his splendid c ollection of butte rflies an d took hi m to a numbe r of his favo rite collec ting local ities. A mong these was King's H ill, where th ey collec ted Argynnis (now Speyeria ) albrigh ti at the typ e locality. La te that season , Garth retur ned to Glacier Park and obtained many lat e - flyin g species o f butterfl ies. An unforgettable memory as th e Garths le ft Glacier Park af ter Labor D ay was of sw arms of Nymphalis j-album alighting in every sunny patch along the highway near Lake McDonald. Following World War II, Garth was able to retu rn to Glacier Park for vacations in l949, l950, and l954. A Park Service ski cabin was made available, making it possible to ex plore vast areas of th e park inaccess ible to to urists, includin g the re mote northwestern lakes, Bowman and Kintla. In l970 and l974, the Garths were joine d for field work at Glacier by Jens W. Knudse n. John's greatest thrill came that last year when Knudsen and Gart h hiked from Lower St. Mar y's Lake to the summit of Cathedral Peaks, where they found Oeneis melissa flying among mountain goats within two feet of the edge of a mile - high precipice. An illustrated booklet on the butterflies of Glacier National Park was projected, but N.P.S. administrative support waned at the time. Today, these unique collections fro m Glacier Nat ional Park are s till to be found at th e Alla n Hancock Foundation , with rep resentat ive series from early years in the Nature Wor kshop at West Glacier, Montana. One hopes that these fascinating su rveys of the distribution and ecology of the butterfly fauna of Glacier National Park will be p ublished by John Garth in another venue; his thorough research in the back country there would provi de a unique historical perspective by which to judge future faunal changes. In the middle of the years of his Glac ier work, John Garth turned his attention to Ariz ona and the butterfly fauna of Grand Canyon National Par k. His field work began in the Park on the Powell Plateau in l946, and was strongly supported by his old fr iend Harold C. Bryant, no w Superintendent of Grand Canyon National Park, as well as Louis Schellbach III, chief naturalist. A well - illustrated book entitled Butterflies of Grand Canyon National Park was published in l950 and is st ill in print. I t combines a scie ntifically detaile d appro ach with a popular writing s tyle tha t engages th e interes t of amateu r natur alist and pro fessional sc ientis t alike. John Garth's latest contribution, California Butterflies , was publishe d in la te l986 w ith J. W. Tilden. It was an outgrowth of their j oint monographic work on the Yosemite butterfly fauna in l962, and Tilden's l965 booklet on the butterflies of the San Francisco Bay region. The two authors were under contract separately to the University of California Press for popul ar works on the northe rn and southern Californi a butterfly fa unas at the ti me (l973) tha t Thomas C. Emmel and John Emmel published The Butterflies of Southern California with the Natura l History Museum of Los Angeles County. The U. C. Press proposed to Garth and Tilden that they combine forces on a new single book as G uide No. 5l in the California Natural History Guides series. The book illustrates m ost of the 235 speci es recorded in the state by Garth and Tilden, usually with one Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 33 northern Californ ia subsp ecies and one from so uthern Ca lifornia. Twenty fu ll - colo r plates by David Mooney add to the impa ct of the text of this excellent introduction to the California butterfly fauna, a book that includes much novel information on the hi story o f butterfly collecting in California that only someone of John Garth's great scholarship and meticulously accurate nature could gather. John S. Garth has had a truly remarkable career, one that has spanned the period from the fabled early lepidopterists of Cali fornia such as Joh n Adams Comstock hi mself to the present -day young students who are them selves contributing so much each year to our knowledge of the western Lepidoptera. A man who intensively studied Lepidoptera before most of those reading this book were born, who travelled t he West in model - T Fords a nd rickety metal traile rs before th e days of Interstate highways, 4 - wheel - drive , and fancy motel s, and who continues to publish his findings and encourage younger workers today, John S. Garth has made an important and lasting impact on the study of western Lepidoptera. Robert Gra nt Wind (19 12 -1975) Robert Wind was one of the most active promoters of the study of Lepidoptera on the West Coast. He was born in San Francis co on June 17, 1912, and died in Guatemala on J uly 4, 1975. He was raised in the community of Berkeley and atte nded the University o f Califo rnia at Berke ley from 1930 to 1933. As a nature counselor of the Boy Scouts of America Camp Wolfeboro in the Sierra Nevada of Stanislaus County in the summer of 1933, he strongly influenced Thomas W. Davies and William A. Hamme r to start bu tterfl y collecting. These men subsequently became leading le pidopter ists in cen tral Cali fornia. Also in 1933, Wind began a new business, Pacific Coast Biological Company, for sales of all types of biological supplies. He i ncluded lists of insect specimens for sale f rom around the world. In 1938, a year aft er marrying Clo Mifflin, he sold his business in order to finance and organize his "Papuan - Australian Expedition." Accompanied by his wife, Wi nd collected from 1938 to 1940 throughout New Guinea, the former Dutch East Indies, and Australia, reportedly returning with a quarter - million spec imens of insects. His health w as affected by these tropica l travels and on his return, he spent so me time rec overing fro m tuberculo sis and then started a ma il - order philatelic business from the sanitorium. He purchased a house in Livermore and continued his phil atelic as w ell as entomo logical s ales, the la tter primari ly from his ex tensive W ind Expedition materials. Wind collected frequently throughout Contra Costa County, twice at Strawberry L ake at Pinecrest w here the new Melitae a leanira daviesi Wind was colle cted, and at Sonora Pass. During the period of 1945 - 1947, Wind actively published on the Indo - Au stralian Lycaenidae (papers co - authored with Harry K. Clench), a s well as on the No rth American Satyridae and N ymphalidae. Eightee n species and subsp ecies were descr ibed in five p apers . From 1950 to 1955, Wind opened and operat ed the Butterfly Tree Park Museum and Gift Shop in Pac ific Grove, associat ed with a motel and a g rove of Monterey P ine where overw intering Monarchs usually stayed. In 1953, the Winds established a business on Fi sherman's Wharf in Monterey. Here they sold natu ral histo ry mater ials, pa rticula rly shells and butte rflies. A large w arehou se had Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 34 to be rented on Cannery Row in Monter ey to have room for all the stocks and this led to t he establishment of Bob Wind's Butterf ly Shop, with floor space of over 20,000 square feet. Insect specimens from all over the world were in st ock here. In 1955, the Winds opened another gift shop called the T rade Winds, also loc ated on the w harf at Monterey. Wind became very well - known nationally because of his promotion of butterflies and insect collecting, and even received awards for his outstanding product developments aiding the growth and welfare of the hobby industry (because of his butterfly - collectin g and mounting kits, which started many a youngster in the field of entomolog y). In May 1959, Wind joined a partnership to open "The Outrigger" restaurant located at 700 Cannery Row on the ocean end of the Old Monterey Cannery overhanging the waters of Monterey Bay. The front end of this cannery was occupied by his wh olesale and retail Butterfly Shop, while the re staurant was capable of seating 600 persons with dining on two levels overlooking the Bay. In October 1960, the Winds opened an even larger Tra dewinds G ift Shop in the Carmel Plaza area. Fin ancial difficu lties in the restaurant business led to a gradual sale of the gift shops and in 1965, Wind moved to Mexico. There, he renewed his activities based on his own collecting of butterflies and insec ts, part icularly in Jalisco and later in the stat e of Chiapas. In 1975, he was moving f rom Chiapas to collect in Guatemala when he died on July 4. Butterfly specimens that Robert Wind collected personally were handled through his wholesale and retail business, a nd today they are found throughout the world in priva te and institutional collections. Some 9,600 insect specimens fr om the tropics were dep osited in the California Academy of Sciences collection before his death. Then in 1986, his collection of western North American butterflies, assembled in the 1930's and 1940's, and num bering 3,841 pinned specimens and including many types o f the species and subspecies he named, was transferred to the California Academy. Taxa named after Robert G. Wind include Incisalia fotis windi Clench, Euphydryas anicia windi Gunder, and an aberration, Plebejus par dalis wi ndi Gunder. A listing of the new taxa described by Wind is given in Arnaud and Davies (1976), but the only new taxon named from Californ ia was Melitaea leanira daviesi Wind, named as a new subspecies in 1947 from Strawberry Lake, Tuolumne County, Califor nia. The Wind Collection included eight specimens of the extin ct Speyeria a diaste at ossa (Edwards) from the Tehachapi Mountains (Arnaud, 1977). Francis Xavier Williams (1882-1967) Williams was b orn in the small to wn of Martinez, n ear the north eastern end of San Francisco Bay, California on August 6, 1882. When Williams was about 5 years old, his family moved to San Francisco. He obtained an A .B. degree in 1903 after attending St. Ignathius College, but rather than entering the priesthood as his family hoped, he went to Stanord University where he studied zoology and graduated in 1908. Another f uture famous lepidopterist, Fordyce Grinnell, was also a student friend there at Stanfor d. Williams became enthusiastic over entomology by his contact with Dr. Edwin C. Van Dyke, at the Ca lifornia Academy of Sciences. However, Williams had started collecting butterflies at Martinez between the ages of 4 and 5 years, and began serious collec ting a nd study even as a small boy. His father made insect boxes for Francis and he had an Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 35 attic room modified as a l aboratory where the young scientist could house his collectio n and carry on his experiments. The family lived close to Lone Mountain in San Franci sco and young Francis could als o collect in the Pres idio ar ea as well. Thus h is ex tensive collections of Sa n Francisco insects were made at a ti me when there was much natural vegetation and open space in areas later covered by roads and buildings. Historically, his material is "the best sample of the now essentially extinct Lepidoptera fauna of the San F rancisco sand dunes" (Powell, 1964). He published a notable paper on the butterflies of San Francisco in 1910, a s well as other observations and notes at l ater times (see Arnaud, 1970, for a complete list of Williams' publications and the insec t taxa described by him). While stil l a student at S tanford, Willia ms joined th e California A cademy of S ciences Expedition to the Galapagos Islands in 1905 - 1906, on which he served as entolomogist. He spent about a year in the archipelago and visited all of the important islands. His import ant publica tions on the Galapagos insect fauna remain classic works to this day, and his collection of Coleoptera there was the larg est ever made in th e Galapagos. In 1910, when Williams published his article "The Butterflies of San Francisco, California" in Entomological News , he listed 4 3 species of butte rflies tha t were know n at that t ime to inh abit or to have inhabited the area, and he stat ed that perhaps not more than 30 species could still be found there in 1910. He also prophetically said, "Before a decade has pa ssed aw ay th ere will be little lef t of the in sect faun a of our cit y, and there fore the imp ortance a nd necessi ty of making local insect li sts while th e fauna is still extant and in an undiluted condition cannot be too much emphasized." His comments on the v egetation and conditions prevailing in San Francisco in 1910 are invaluable today and in striking contrast to what we now see in the area. In 1913, Williams went to Harvard to obtain hi s D.Sc. (conferred in 1915) under Professor William Morton Wheeler, and he subsequently went to Hawaii to work on insect problems in the sugar cane plantations. During the next five years, he traveled to the Philippines in search of parasites an d predators, and made extensive no tes on the hab its of the bees and wasps of th e Hawaiian Islands. He continued economically oriented research throughout the tropic s until 1948, when he retired and moved to Danville, C al ifornia, near the base of Mt. Diablo and east of San Francisco Bay. E ventually, the Williams moved to La Mesa, near San Diego, C alifornia, where Williams became as sociated closely w ith the San Diego Museu m of Natural History an d continued his study of w asps. While Willia ms greatest fame is as a hymenopterist, h e was an ext remely broadly based researcher whose early writings on butterflies di d much to record the passing scene in the San Francisco area of California (Zimmerman, 1969; Arnaud, 1970). William Ba rnes Barnes was born in Decatar, Illinois, on September 3, 1860, and died on May 1, 1930 at the age of 70. At this t ime, Dr. Barnes owned the la rgest and most important private collection of North Amer ican Lep idoptera. Residing in Decat ar, Illino is, for mos t of his l ife, he was also one of the fore most surgeo ns in th at state. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 36 After graduation from Harvard University (1883) and Harvard Medical School (1886), and a post - graduate medical study period in Germany, Barnes returned to I llinois to begin his career as a surgeon and to carry on his avocational researches in N orth Americ an Lepidoptera. In the course of this work, he assembled one of the outstanding world - class collec tions of American Lepidoptera. He hired the best possible curatorial staff, i ncluding J. H. McDunnough, F. H. Lindsey, and S. H. Benjamin, all trained entomologists of outstanding ability. His collection was housed in a separate buildi ng from his home, with fireproof construction, and oak cabinets containing some 1,200 drawers with over 2,000 additi onal Schmit boxes arranged on shelves along the wall. The type specimens, i ncluding the holotypes, paratypes, and the li ke, amounted to nearly 7,000 specimens. The entire collection numb ered over a half million specimens. As far as California lepidopterology is concerned, Barnes' chief goal was to obtai n some of the Boisduval types of California sp ecies. Secondari ly, he purchased Califo rnia specimens from Ximena MacGlashan and E. J. Newcomer. He also publ ished the copiously illustrated se ries of volumes, "Contribu tions to the N atural Histo ry of the North A meric an Lepidoptera," which included the taxonomic r esearch by Barnes and his collabo rators and co nsisted of e xtensive descriptive and revisionary papers, including California ma terial. The Barnes collection was purchased by the United States government t hrough an act of Congress and was depos ited in the U.S. National Museum in Washington in 1930 (Engelhardt, 1930; Schaus, Busck, and Heinrich, 1930). William Ne lson Bur dick (1887 -) Burdick was born in Denver , Colorado in 1877, the son of a well - known physician. He graduated from the University of Cali fornia with a degree in ent omology which he pursued as an avocation throughout his life. He authored a number of scientific papers on N orth American butterflies and bequeathed his large collection of over 20,000 specimens to the University of Colorado Museum at Boulder (Anonymous, 1962). Erich Carl Walter ( 1903 -1990) Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 37 Christopher Henne (__ __ -______) Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 38 Edwin Ray Hulbirt (1886 -1965) Born in Pennsylvania on April 9, 1886, Hulbirt died in Glendale, California, on March 12, 1965. A fter rece iving h is college degree in chemist ry from Princeton Universi ty, he taug ht in Oregon and then Azusa, California, helping to organize the Citrus Junior College and serving as its first dean from 1915 to 1918. His chem ical research led to his appointment as Chief Chemist for a company that eventually was absorbed by the American Cyanimid Company, in which he served as Chief Chemist from 1920 to 1941 and Administrative Assistant from 1941 to 1947, when he retired. He had a very stro ng avocatio nal inte rest in ento mology as we ll as mounta in climbing . Between 1909 and 1923, he climbed many of the great peaks in the western United States, including (in California) Mt. Shasta, Mt. San Antoni o (Mt. Baldy), Mt. San Jacinto, Mt. Hoffm an, Mt. San Antonio, Mt. Whitney, and Mammoth Peak. He collected butterflies throughout the state of California and particularly specialized in the Hesperiidae and Lycaenidae. Two butterflies w ere named for him, Hesper ia comma hulberti Lindsay (1939) and Lycaena dorcas hulberti Field (1936). His collection of 23,000 Lepidoptera was given to Citrus Colle ge in Azusa, California, in 1963 (Comstock, 1965). After Hulbirt gave his worldwide collection of Lepidoptera to Citrus College in January 1963, the school, fa ced with a shortage o f storage and exhibit sp ace, transferred the entire collection in late 1972 to the Natural History Museum of Los Angeles Count y as a permanent loan. This collection then consisted of 19,340 specimens (Donahue, 1975). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 45 Butterfly farms are becoming r elatively common around the world today, from scores initiate d on small to larg e scales i n Europe to major recent en terprise s in Japan, Australia , Malaysia, the Philippin es, and several huge att ractions in the U nited States. But Ch arles Fayett McGlashan and his daug hter Ximena star ted such a farm in Truckee, Calif ornia, at the turn o f the century and made quite a success of it, reaping great national publicity at that time, long before these rela tively la te "new comers." Charles McGlashan was born in Wisconsin in 18 47. After Charles' mother died two years later, his father eventually decided to gather up his family in 1851, load them into a covered wagon and head for California. Consequently, Charles grew up in the rugged mining and timber country of the Sierra Nevada and by 1872, he found himself a job as superintendent of the school system of Truckee, a Sierran town on the ro ad between Sacramento and what was later to be come Reno, Nevada. Th e Central Pacif ic Railroad had arrived just th ree years earli er and had helped to transform Truckee from a stopover on the great east - west emigrant wagon trail in to a bustling terminus on the new transcontinental railroad run. McGlashan was an ambitious young man who tried his hand at many occupations, including tea ching school, studying law, collecting butterflies and moths, making historical notes on regional lore, and serving as a local corres pondent for the Sacramento Daily Record -Union . By 1875, he was not only a l awyer in Tru ckee, bu t also editor of the local newspaper, The Truckee Republican . He beca me famous acr oss the United S tates in 1879, when he published his book, History of the Donner Party . This melodramatic and grisly account was based on actual interviews with the survivors of the Donner Party still living at that time. It told the true stor y of an emig rant party th at started la te in the sum mer to cross the Sierras. They were caught by a snowstorm near Truckee, and the survivors had to resort to cannibalism on the bodie s of their f riends and relativ es to live un til rescu ed. In 1893, McGlashan built a museum containing collectio ns of baskets made by local Indians, Donner Party relics that he had excavated, and butterflies and moths that he had col lected in the local area. McGlas han had first learned how to collec t and rear L epidoptera from an English prospector named Oldfield who was an insect collector. Not long afterwards, he met Henry Edwards, an ardent entomologist who worked the mining - ca mp entert ainment c ircuit as an actor. For the next dozen years, he shipped Edwards many of the moths that he was able to rea r and he also corresponded with other promi nent American entomologists of the day. Eventu ally, he was able to amass a collection of over 20,000 speci mens, part of which survive today in the Nevada County Courthous e, in Nevada City , California, not far from Truckee. In 1912, McGlashan and his seventh daughter, Ximena, hit upon the i dea of building a butterfly farm. Two decades ear lier in England, a young lepidopterist named Leonard W . Newman had started a successful butterfly fa rm in England, and it is possible that McGl ashan had heard about this f arm. The ear ly twentie th century in America w as an appropr iate time to b e starting new enterprises, and McGlashan's farm turned out to be a huge s uccess. During that summer of 1912, Ximena was 18 years old and had just gr aduated f rom San Jose Normal Sch ool. McGlashan casually remarked to her that he thought she could earn more money catching butterflies than teaching schoo l. He had received an offer f rom Dr. William Barnes, the wealthy midwestern collector, to purchase any Lepidoptera that he could collect and ship, but Charles was far t oo busy with his law practice to get involved . Ximena was quick to realize the p ossibilities of this o ffer Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 46 and within two weeks, after mastering the skills of collecting and rearing, she had shipped Barnes about 1,500 specimens for which he paid her $75. By the end of that first su mmer, Ximena had shipped over 10,000 specimen s, being paid at a flat rate of five cent s a specimen. This averag ed out to about $50 a week, which was certainl y more money than she could have earned teaching school. After a loca l new spaper article was published o n her activities, the Hearst new spaper chain p icked up the story of her exot ic and unusually profitable business. Soon, she was receiving hundreds of letters from people who wanted to imit ate her success. Thu s in Septem ber 1913, she and her father began publishing The Butterfly F armer: A Monthly Ma gazine fo r Amate ur Entom ologists . This magazine was full of practical information, and consisted essentially of a comprehensive correspondence course in entomology. Suggestions of books to read, equipment needed for collecting (and how to make it or where to buy it), how to care for eggs, larvae, pupae, and adults, how to mount spe cimens, how to sugar for moths, etc., were all contained in the monthly issues. In September 1914, Ximena enrolled as a student of entomology at the University of California at Berkeley and transferred to Stanford a year later, where she got a degree in botany and entomology. Her success in business continued until World War I. Then , marriage to a p rofessor of milita ry science (who had to move to Fort Leavenworth, Kansas, to train enlisted men) intervened, and she never went back to farming butterfl ies. This summa ry of her life is dra wn from Oppew all (1979), who intervie wed Ximena McGlashan Howard at B erkeley for her definitive biography of this fascinating personality and her father. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 47 Albert and Amy C arter The next se rious butte rfly farme rs in Califo rnia had more similarit ies to presen t - day butterfly farms and parks, in that they opened the ir rearing facilities to the public and had a specially landscaped, hillside insect zoo where the public could roam at l arge for free ev ery Sunday. Albert and A my Carter b egan thei r butterfly farm as a get - rich - quick scheme in 1929, and called their enterp rise "Bu tterfly Park." Th e first issu e of their magazine, Bu tterfly Park Nature Club News , was s ent out in March 1929 , and it appeared monthly through October, whe n the great 1929 stockmarket crash began putting a dent in all business enterprises. The Carters' publication then became bimonthly, it became quarterly in 193 1, and petered out sometime in early 1932. Their magazine contained essentia lly the sa me kind of i nformation as Ximena M cGlashan's Butterfly Farm er , and the Club managed to attract about 500 to 600 subscribers. The Carters used their reared butterflies to make into s uch objects as book ends, trays, and jewelry. T hese artistic creations were sold at the Butterfly Park, located in Roscoe, now called the "Shadow Hills" section of Sunland, in the northern end of the San Fernando Valley in southern California. Lloyd M. Martin, who was to beco me Associate C u rator of L epidoptera at the Natu ral Histo ry Museum o f Los Angeles County, worked for the Carters for five years fro m 1930 to 1935, the last year being the date that the Park was closed down. Up to that time, howev er, as many as 100 visitors a Sunday came to wander freely in the 100 - sq uare - foot, spec ially landscaped , screened - in h illside in sect zoo where, as the Carte rs said, "16,000 c aterpillars are feed ing on foodplan ts planted especial ly for them." C ascading w aterfal ls, inviti ng picnic tables and stro ll ing peacoc ks offered touch es which we see today, 60 years later, in so me of the bigger butterfly parks in the United States and elsewhere in the world (Oppewall, 1979). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 48 VI. THE MO DERN AGE OF L EPIDOPTERA ST UDY: RECENT LEPI DOPTERI STS IN CALI FORNIA (1940 - Present) William Ho vanitz (1915 -1977) Born in Chicago on November 6, 1915, William Hovanitz died on September 14, 1977. At the age of th ree, he moved with h is family to the San Francisco B ay area of Cali fornia where his interest in n ature, especi ally butterflies, s tarted. E ven in high schoo l while he was working as a delivery boy for a San Francisco drug store, he t ook time to collect specimens while making his deliveries around the city. These included var ious Plebejus blues that are now rare and threatened , and Glaucopsyche xerces , that is now extinct. He obtained his B.S. degree in entomology fro m the University of California, Berkeley, and in 1938 began advanced study immediately in the graduate program at th e Califo rnia I nstitute o f Technolo gy under N obel - Laureate Th omas Hunt Morgan, where he received his Ph.D. in Genetics in 1943. Because Morgan died during his graduate work, Hovanitz finished under the eminent geneticist A. H. Stur tevant. Hovanitz spent the next thr ee years (1942 - 1945) w orking for the National Research Council in Sou th Amer ica, Michigan, and Florida. He then accepted academic appointments at the Laboratory of Vertebrate Zoology at the University of Michigan, Wayne University at Detroit, an d finally the University of San Francis co. At the latte r instituti on, he serve d as A ssociate Editor of the Wasmann Journal of Biology . While teaching summer s essions a t the Un iversity o f Californ ia at San ta Barbara in 1949, he met and married his wife, Barb ara. In 1 955, he re turned to Cal Te ch with so me time sp ent at U.C.L.A., but finally settl ed into an a cademic positio n as Professo r of Biology at the Californ ia State University in Los Angeles in 1956. He r esigned from that faculty in 1975, and in e arly 1977, the Hovanitz's moved to Santa Barbara whe re they planned to build a hom e and allow Bill to devote his full energies to research. His untimely death cut those expectations short, particularly unfortunate because of the large body of unpublished materia l that Hovanitz had ready for analysis. Hovanitz published 131 papers and other publications during his lifetime. Some o f his most impressive and significant works were his 1943 pape r on the variation on Argynnis c allippe in California, and his 1940 paper explaining color variation of Oeneis chryxus in the Sierr as, in particular the occu rrence of the dark - colored subspeci es stanislaus and the light - col or ed subspecies ivall da as related to the distribut ion of their alp ine backgrou nd substrates. One of his earliest papers, w ritten as a sophomore in co llege, described the geo graphic variation in the lycaenid butterfly, Plebe jus icariodes in terms of its relat ionship to a Rassenkreis ("c ircle of races"). He was perhaps most f amous for his work on Colias butterflies, which included several classic papers such as hi s 1948 treatise demonstr ating differences in diur nal activity of genetical ly differen t orange and white female s with Colias eurytheme in the Imperial V alley of Ca lifornia (explaine d in terms of a daptation to d iurnally cy clic environ mental fac tors). He a lso looked at th e influence of compl ex enviro nmental selectiv e forces in producing c olor varia tion in the female dimorphis m of Colias species across the cont inent (Hova nitz, 1950). His classic studies on geographic variation in Euphydryas chalcedona (Hovanitz, 1943, 1952) revealed the tip of a scientif ic iceberg of inc redible d imensions and complex ity in this g roup, wh ose details are s till being explored by dozens of lepidopt erists and evolutionary biologists today. He also contributed very signific antly to butte rfly bioge ography with th e first compre hensive rev iew of the Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 49 zoogeography of butterflies in the New World, publis hed in 1958 by the American Association for the Advan cement of S cience. Th is majo r paper desc ribed the ma jor fe atures of lat itudinal an d altitudin al distrib ution of the fa milies an d major group s of butte rflies in the N earctic and th e Neotropics. One of Hovanitz's most enduring legacies, of cou rse, was the establishment of the Jou rnal of Research on the Lepidoptera i n 1963. This effort was not without controversy, as it was viewed by some as competition with the existing Journa l of the Lepidopterists' Societ y . But as Mattoni (1979) pointed out in his definitive biography of William Hovanitz, "a clear niche developed for both to th e lasting b enefit of a ll lepidop terists." After 40 years of collecting and studying butterflies before his untimely death in 1977, Hovanitz had desposed of parts of his material to the California Academy of Sciences. The William H ovanitz ma terial sti ll in his fam ily's p ossession at his dea th num bered almost 24,000 specimens, and this material was given to the A llyn Museum of Entomology (part of the Florida State Collection of Arthropods) in 1981 (Miller, 1983). Several years later, the more than 4,400 Colias and other Pieridae groups from the Ne arctic and Pa learctic wer e transferred to the Califo rnia Academy of Sciences collection. In 1992, the Pacific Slope Section of the Lepidopterists' Society posthumously conferred upon William Hovani tz the John Adams Comstock Award in recognition of his manifold contributions to the study of California butterflies and Lepidoptera in general (Emmel, 19 92). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 50 Lloyd Milo Martin (1912-19xx) Lloyd Milo Martin was born on a homestead near Bayard, Nebraska, on May 24, 1912. After the death of his fat her in a farm accident four years lat er, he was raised by his grandparent s in Steele City, N ebraska. T rips back to the f arm be came his f irst colle c ting forays as he helped a cousin collect butterflies for her collection. In December 1921, Martin moved to Long Beach, California, with his grandparents, and his collecting activities increased. In his own words, "Many species o f butterfl ies, so diffe rent from th ose of th e farm in Nebraska, w ere caught i n Californ ia; this was the beginning of a real i nterest in insects. It was my ambition to get a college degree and, some day, to work in a museum. The depression of the early 1930's put an end to plan s for college. After graduation from high school in 1929, I worked for Hal Newcomb in South Pasadena, doi ng butterfly art work. This work went o n for about a year. From May 1930 to Oct ober 1935, I worked at Butterfly Park, near R oscoe (now Sun Valley), for Albert C arter. [T his comme rcial e ffort by Carter to build a showplace of butterfl ies ultimately failed.] During the summers of 1931 and 1932 I was in th e field , collec ting but terflies fo r Jeane D. Gun der of Pasade na. I mad e two t rips to the Catalina Mts. of southern Arizona in the spring of 1931; later in the same year I collected at Pyramid Lak e, Nev., in the Cas cade Mts. of n orthern Cali fornia, includ ing Mt. Shasta. The summer of 1932 was spent in the field in southern California." On February 19, 1936, Lloyd Martin started working f or John Comstock at the Natural History Museum of Los Angeles County as a Student Assistant in E ntomology. Like Comstock, he donated his personal collection to the Museum. In 1941 he was promoted to As sistant Preparator, then to Curatorial Assistant in 1943, and on July 1, 1944, to Assistant C urator of Entomology. In 1952 he was promoted again to Associate Curator, in charge of all Lepidoptera at the Museum (some 250,000 specimens at that time). He held this position until his retirement on July 31, 1969, at which time he was given the title of Curator E meritus. He continued as a Research Associate of the Museum, operating from his home in Prescott, A rizona, where he avidly continued his research es on the moths of that fascinating state until his death on _____, 19__.. During his three decad es at the Museum, Lloy d Martin carried out th e usual curatorial duties of collecting, classifying, and preparing insects for the museum collection and f or exhibitio n, as well as identify ing inse cts for sc ientific go vernment a gencies and the general public. His greatest source of pleasure, though, was encouraging young collectors through the Lorquin Entomological Society meetings and field trips, and through answering thousands upon thousands of questio ns from th em, espec ially every S aturday up on t he "third floor" of the Museum at Exposition Park. Under Comstock and then M artin, the holdings of the Entomology Section of the museum greatly i ncreased in size and diversit y. Comstock and Martin were both active collectors and outgoing individuals who attracted the volunteer cooperation of many amateur collectors, old and young, in the Los Angeles a rea. This helped to accelerate the growth of the collection . Lloyd Martin wrote a major paper o n the butterfly fauna of t he Huntington Lake region in Californ ia, published an annotat ed check list of North A merican L epidoptera in the Natura l History Museum of Los Angeles County, described several new butterflies inc luding Papilio indra fordi Comstock & Martin and Euchloe hyantis andrewsi Martin, and wrote a widely - noted article for the general public, "Spectrum on Wings," for th e Arizona Highways magazine (April 1951 issue), Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 51 describing the beauty of Lepidoptera found in the Southwest. He intensified his personal field work in Arizona together with Fred S. Truxal, and the museum rapidly accumulated an outstanding collection of butterflies, moths a nd other groups from the littl e- known insect fauna of that state. Martin recounted in an a utobiograp hical ske tch that, "My re al interest was trying to find o ut how insects could survive in the se mi - arid regions an d deserts of the Southwest. I was fascina ted by their adaptations to the environment, the changes of color patter ns to blend in with the desert, the vast numbers of species of insect s that are noctu rnal in hab it, and the co mparatively few diurna ls, due to the inclement conditions in which they live." His photographs and vivi d d escriptio ns of these studies at many national and regional meetings of the international Lepidopterists' Society, the Southern California Academy of Sciences, and the Lorqui n Entomological Society, brought to thousands of budding naturalists and professional biologists the richness of the natural history of the American Southwest. Despite all t he preceding acco mplishments, Llo yd Martin alw ays maintained h is strongest enthusiasms for helping beginning students of Lepidoptera. H is finest legacies are the legions of southern California entomologists and biologists who have been profoundly influenced to go into serious amateur or professional work with Lepidoptera because of his dedicated example and constant encouragement. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 52 Charles L. Remington Charles L. Remington was one of t he two co - founders of the Lepidopterists' Society in 1947 and has served in this Society and in the development of the science of lep idopterology in the United S tates for more than four decades. His researches in the biolo gy of west ern Lepidoptera, particularly in Colorado, Texas, northern Mexico, and the Channel Islands of California, have made important contributions to our knowledge of the evolution, ecology, biogeogr aphy, and life histories of our western species. His editorsh ip of the Journal of the L epidopterists' So ciety during its formulative years included not only remarkable expansion of its scientific and amateur scope, but also the careful nurturing of many of our western lepidopterist authors and researchers. Charles Lee Remington was born i n Reedville, Virginia, in 1922 and he grew up i n St. Louis, Missouri. He pursued his undergraduate w ork at Principia College in Illinois, receiving his B.S. degree in 1943. He migrated eastwar d to Cambridge, Massachusetts , where he earned his Masters in 1947, and his Ph.D. at Har vard University in 1948. During those graduate school ye ars at Harvard, he and Harry K. Clench, who knew each other by previous correspondence due to thei r mutual in terests, go t together and dec id ed to promote the formation of a union or soc iety of lepidopterists that would encourage the scientific study of Lepidoptera. This joint aim was to be accomplished through distri buting a periodical on that insect order and through facilitating the exchang e of specimens and ideas by both the pro fessional worker and the interested a mateur. This effo rt formally st arted on Marc h 24, 194 7, with a lette r to the w ide circle o f friends that each man had in th e field. The dues were a modest $1.00. Char les Remington was the first Treasurer in this founding year of the Lepidopterists' Society. Th e Journal began that year as an inexpensively mimeographed publication. It graduated to photolithography when revenue and membership grew enough to warrant the change. From there, it eclosed to regular pri nting. As Harry Clench has said in the Society's commemorative volume, Charlie Remington knew what he wanted and how to go about getting it. L epidopterists, evidently, agreed with him. This Lepidopterists' Society, which he largely shaped, has grown and prospered to major size and influence in the fie ld today. In the beginning, no meetings or elected officials were contemplated. By 1950 , however, memb ers realized that forma l organiz ation wo uld be in the best interests o f the growing group and on July 31, 1950, Char les Remington appointed an organi zat ion committ ee, naming Cyril F. dos Passos as chairman, with the charge to ap point temporary officers and develop the Constitution and By - Laws. Th e present for m of the Soc iety resulte d from this committee 's efforts, an d the first an nual meeting o f the Socie ty was held 29 - 30 December 1950 at the American Museum o f Natural H istory. In S eptembe r 1954, the first Pa cific Slope Section M eeting o f the Lepidopterists' S ociety was held in Califo rnia at the Academy of Science in San Franci sco. Pacific Slope Section annual me etings have been held each year sinc e l954 and o ccasionally these hav e been combined with the national meetings of the Society (l 962 Santa Barbara, 1970 Carson City, 1974 Los Angeles, 1976 Boulder, 1982 Laramie, 1987 Berkeley, 1989 Albuquerque, 1991 Tucson, 1993 Fort Collins). Though an east ern member, Remington participated on the programs of the 1965, 1974, and 1988 Pacific Slope meetings, and attended others during the 1960s. Publications of this Society were also heavily influenced by Charles Rem ington. The Society's first publication, which began in 1947, was known as The Lepi dopterists' New s . Under Remington' s editorsh ip, this o rgan evolv ed from an initial mimeograp hed format to a Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 53 photolith ographed fo rmat and finally to a le tter pre ss format from Volu me 6 onw ard. Wi th Volume 13 in 1959, at Remington's urging, the name of the Society's major publication was changed to the Journal of the Lepidopterists' Society , and the Society initiated a second, more frequent and informal pu blication, The News of the Lepidopterists' So ciety , which was designed to include matters o f more transient in terest. Charles Remington serve d as Edito r - in- Chief of the Journa l and its predecessors from 1947 to 1963. It is also of great interes t to note th at the fi rst Annua l Field S ummary was b egun at the same time by C. L. Remington and Harry K. Clench, founders of the Lepidopterists' Society. North Amer ica was div ided into e ight zon es in their initial e fforts, and by the time of the second annual (1948) summary, each had a corresponding "zone coordinator." Thus, from the very first year of the Societ y (1947), individual members were encouraged by Remington and his c olleagues to contribute to the Summary and to keep careful fiel d notes during each collecting season. In the first year, there were 59 contributors and th is has increased to several hundred today. Charles Remington also started the Library of the Lepidopterists' Society, which today is housed at th e Na tural Hist ory Museum of Los A ngeles C ounty in C alifornia but was started a t Yale University . This evolved from the publications being received by Remington in exchange for copies of T he Lepidopterists' N ews . The Society Library contains comple te or nearl y complete s ets of some forty periodicals devoted in whole or in part to entomology, and incompl ete sets of many more. In addition, there are many reprints and a large collection of books which have been donated over the years by authors and members. From the beginning, the policy ha s been to make this material available on loan to members upon request. In addition to his contributions of ine stimable value to the founding and evolution of the Lepidopterists' Society, Charles Remington has had a r emarkably diverse research career. The range of his specialties extend f rom evolutionary genetics to speciation theory and experimental and natural hybridization in animals and plants. He ha s delved into island biology and biogeography, mimicry, insect - plant coevolution, the systematics of Lepidoptera, paleoentomology, and even cytosystematics of insects. Collaborating with K. Maeki, he pioneered the study of chromosomes in the North American Rhopalocera, uncovering man y fascinating cytotaxonomic situations, especially in the Megathymidae. In thi s latter family, he pursued profitable collaberations with Don B. and Viola Stallings, the Turners, and Hugh Avery Freeman throughout the southwestern United States and northern Mex ico. His field research has taken h im to Australia, Iceland, England, Norway, Italy, Mexico, Canada, the off shore islands of California, the Hawaiian Island s, the Marshal l Islands, Nauru, G uadalcanal, the New Hebrides, New Caledonia, New Zealand, Fiji, Tonga, Moorea, Bora Bora, Tubuai , Layte, Hokkaido, Ecuador, the Galapagos Islands, Peru, Brazil, Chile, Venezuela, and 49 states in the U.S.A. He has collected Lepidopte ra on all eight o f the Cali fornia Chan nel Island s, parti cularly on San Clemen te Island in August 1968 and December 1981 and extensively on Santa Cruz Island in all the spring months except February and in July, August and Oct ober, between 1967 and 1974. He deployed his son Eric there for about three months in 1970, and many moth records from Santa Cruz in the California I nsect Survey come from his efforts. C harles was one of the organizers o f the 1978 Channel Islands Symposium a t Santa Barbara and presented an overview of the insects in a plenary session paper. H e likew ise participated in the 1981 E ntomological Scoc iety of America symposium on Channel Isl ands insects. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 54 Despite his intensive research progra m and field travel, he found time to b e an outstanding teacher at Yale University from the onset of his appointment there in 1949. He rose through the academic ranks to the highest position as Professor in the Department of Biology, Curator in Entomology of the Peabody Museum, and Professor of Forest Entomology and Mu seology in the School of Forestry and Environmental Studies. Among his many academic honors, he was a Sheldon Traveling Fellow at Harvard University in 1947, a John Simon Guggenheim Fellow at Oxford University in 1 958 - 59, was named an honorary Li fe Member of the Lepido pterists' Society in 196 6, was named a R esearch Fellow in Entomology for C.S.I.R.O. in Australia in 1976, a Research Fellow in Ecology at Campinas University in Brazil in 1981, a Researc h Fellow in Entomology at the University of California, Berkeley, in 1981, and has been an invited lecturer at many n ational and international meetings. He h as been Secretary (195 5 - 59) and Trustee (1962 - 63) for the Rocky Mountain Biological Laboratory in Gothic, Colorado, where he has acti vely pursued field research for more than 20 summers, including work there with Dr. Gary Bernard of Yale on the color vision of butterflies. He has served on the advisory boards of the Connec ticut Audubon Society, the IUCN, the Connecticut State Geological and Natural History Survey, and as Counselor of the Xerces Society. He w as name d Honorary Life Member of the Lepidopterists' Society in 1966. Dr. Remington has contributed to the development of lepidopterology in California and the western United States not only through his publicati ons and his participation in meetings and as a visiting r esearch f ellow at w estern ins titutions, but also through his personal enco uragement of scores of western lepidopterists who today are highly active in the Society and in Lepidoptera research. His friends and colleagues have long stood in awe of the indefati gable energy, boundless interest s, and incred ible bre adth o f knowledge of this "Founding Father" of the Lepidopterists' Society, who has brought so much favorable worldwide attention, from sophisticated scientific research to greatly expanded amateur interest, to this fascinating group of insects (Emmel, 198 8). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 61 informat ion on Californ ia butter flies. T his is truly a collabo rative and cooperative p roduct. Its aim to prese nt as comple te a picture a s possible of th e California b utterfly f auna at the c lose of the Twentieth Century : its b iodiversi ty, present and past distrib ution, h istorical c hanges in popula tion sizes, ecology, life history, host plants, chromosome numbers, and overall biology. Our success in reaching such a goal wi ll be judged by you, the re ader. Doubtless, much remains to be lear ned about the bu tterflies o f Califo rnia, and it is the hop e of the a uthors that this book w ill encour age the presen t naturalis ts of the st ate and else where to w ork for the p reservat ion of the sta te's remaining wild areas so that future generations may enjoy what we can still see and study today. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 62 Chapter 2 THE GEOLO GICAL LAN DSCAPE OF CALIFO RNIA In this brief chapter, w e shall look a t the geolog ical history of Califo rnia and its in fluence on the vegetation and butterflies of the state . Our understan ding of the div ersity of th e Californi a flora, and the rich butterfly fauna, depends to a considerable extent on an appreciation of the geological forces at work and the timescale involved in creating the present - day physiographic appearance o f Cal ifornia . Geology has dire ctly i nfluenced c limate, w hich in turn h as dramat ically affected p lant and b utterfly communiti es in the s tate. The following d iscussion is summa rized principally from the excellent essay by Daniel L. Axelrod in Munz and Keck (1963). Early in the Cenozoic Era, which began approximately 63 million years ago, the vegetation across North America wa s composed of three geof loras. A geoflora is a ma jor vegeta tion un it that has managed to maintain its essential identit y through time and space. Throughout the southern half of the continent spread the broad - le aved evergreen Neotropical- Tertiary G eoflora . The northern half of the continent was occupi ed by the temperate Arcto - Tertiary G eoflora , composed of mixed deciduous hardwoods and conif ers. Between these two geofloras, centered in the area of southwestern North America, the Madro- Tertiary G eoflora was making its init ial appearance. Compose d notably of the sclerophyllous ("har d - leaved") a nd microp hyllous ("small- leav ed") plants, the Madro - Tertiary Geoflora contained drought-adapted plants. During the first several epochs of t he Cenozoic Era, the Neotropical - Tert iary Geo flora migrated northward as far as southeastern A laska on the coa st and close to the Canad ian border in the inter ior. Here it came in to contac t with the A rcto- Tertia ry Geofl ora wh ich occupied the higher, cooler la titudes. B road - leaved ever greens of trop ical families, includ ing figs and s abal palms, occurred throughout the area -- species tha t are now at least sub tropical in charac teristics and occu r in region s where ra infall is 80 or more inches annually and where climate is uniformly warm throughout the year. Thus no desert fl ora yet existed in the Southwest, nor was t here any high mountain range in the in terior to fos ter an alpin e flora. The d istribut ion of this ne otropica l forest gradually was restric ted towards the south and to the coasts as a cooler and dri er climate predominated, and no plants in California today represent direct descendants of this humid subtropical forest. However, a number of California genera and species were derive d from that secondarily, as Axelrod ( in Munz and Keck, 1963) points out. The weste rn part of the Arcto-Te rtiary Geo flora bec ame adap ted to a ne w climate, one typified by winter rain and summer drought. The major communities of this geoflora developed chiefly during the later Pliocene, when more diverse climates developed over the Southwest in response to important topographic changes: c hiefly, the elevation of the Sierra Nevada, the Coast Ranges, and the Transv erse and Peninsular R anges. All of the se mounta ins are pa rt of th e great W estern Co rdilleras b elt alo ng the ou ter continen tal edge o f North A merica (the Easte rn Cordil lera is the R ocky Mountain s ystem, alo ng the inner edge of the old North American continent). The Western Cordilleras began forming in the Jurassic Period (about 180 million years ago) of the Mesozoic and continued right on into the Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 63 Miocene (about 10 million years ago). At this t ime, the Sierra Nevadas and the Cascades were initially raised and folded from older Mesozoic conglomerates and coarse sands that had w ashed into the Pacific Ocean from the centr al continent area. Intense fo lding, uplift, and severe faulting followed, especially between 100 and 50 mil lion years ago. Summits in the Sierras eventually reached close to 15,000 feet above sea level. The Coast Ranges have a less pronounced, more rounded relief that is generally below 3,300 feet elevation, but some peaks rise above 6,000 feet or even up to 11,500 feet (3,506 meters) on Mt. San Gorgonio in t he eastern Transverse Ranges. These young ran ges arose in two phases. In the mid - Miocene (about 20 million years ago), earth movements thrust up the Early Cenozoic (60 million year old) layers of sediments that had been washed out along the continental shelf by erosion from the then - exi sting marg in of the North A merican c ontinen t to the ea st. The se upfo lded layers were eroded subsequently almost to a plain, and were covered by new layers of younger sedimentary rocks washed down from the east . Then a second phase of mountain - building began in the Quaternary Period, possibly one million years ago, and has continued to the pr esent as the central Coast Ranges and Transverse Ranges are still being built up, deformed, and eroded today. All these va st mountain ranges alon g coastal an d interior Cali fornia alt ered the cli mate substant ially, w ringing much of the moistu re from the p revailing westerl y winds as th e air masses flowed up and over the mountain ridges, cooling at the higher altitudes and consequently precipitating their water load previously picked up over the Pacific Ocean. Thus the orogeni c (mountain- building ) episode s in the v icinity of California initia ted a wi despread cl imatic cha nge to cooler and drier conditions. Van Devender (1977) presents evidence that the shi ft to non - pluvi al climates in the Southwest occurred as recently as around 8,000 year s ago. This involved a synchronous change from woodland to desert or grassland across the Chihuahuan, Sonoran, and Moja ve des ert s bec ause of a shift to th e north of the A leutian low and the w inter storm track, w hich result ed in drastic ally reduced winter p recipitation in thes e areas. The existence of continental gl aciers to the north had modified g eneral ci rculation pa tte rns and caused the Pluvi al (wet) southw estern climates during the Pliocene and Pleistocene (2 million to 10,000 years ago). With a shift from gla cial to non- glacial climates, the ch anges in pluvial clim ates would lag behind chang es in glacial climates as long as the glaciers existed. The rel atively xeric woodland found between 11,000 and 8,000 years ago in the Southwest may have occurred in s uch a lag period. Finally, when heavy winter precipitation diminished in the Southwest and post - pluvial conditions dried the playa lakes, the formation of desert and grassland communities followed. Regardless of the exac t time table, the new mountain rang es caused the Madro - Tertiary Geoflora, n amed fo r the Sierra Madre of nor thern Mex ico where m any of its relics n ow survive , to origina te in south western N orth Ameri ca. Here, small - leaved, drought -deciduous, sclerophyllous plants were apparently derived from the more humid tropical groups of the Neotropical- Tertiary Geoflora which were becoming adapted to the expanding dry climate. Some members of the Madro - Tertiary Geoflora, however, apparently are d erived member s of Arcto - Tertiary groups which also were developing in response to incr easing aridity during the middle and later T ertiary times. P lant commu nities such as Oa k- Conifer Woodland, Chaparral, Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 64 Thornscrub, D esert Grassland, and Sage, as w ell as part of the Desert flora, app eared from the middle Eocene through the following epochs. By the middle Pleistocen e, dry climates were rapidly expand ing across the area, and Oak - Conifer Woodland and Chaparral vanished from the lowlands of the present central and southern desert regio ns, as well as from the forest of t he northern portions. In the late Pliocene a nd Pleistocene (1 mi llion years ago), the rapidly elevating Cascades, Sierra Nev ada, and the Peninsular R anges created a drier cl imate on their interior ( rain shadow) sides, and a desert flora of remarkably drought - resistant p lants came into existence. T he new desert spec ies had been derived principally from the geofloras that had dom inated the area earlier in the Pliocene (2 - 1 million years ago). The higher and colder Great Basin Desert and northern Mojave Desert received floral co mponents f rom the survivin g Arcto- Tertiary Geoflora. The lowe r and hotter Sonoran and southern Mojave Deser ts obtained species from the Madro - Terti ary Geoflora (Axel rod, in Munz and Keck, 1963). The rainfall during this span of time changed from possibly 80 inches or more in the early Eocene, for much of the West, to about 12 to 25 inches during the Miocene, on the western border of the Mojave Desert. A flora new to thi s region developed as a result. Summer dryness throughout what is now California became important, and the sclerophyllous types of woody plants increased in number. This drought - resistant, small - leaved a nd deciduous flora stood in strong contrast to the p lants typ ical of the earl y Te rtiary floras. By the Pliocene, the forest and woodland had to move towards the coast or into the mou ntains, and the lower elevations of what is now desert became domin ated by scrub and grassla nd. Desert species evolved from plants already present, t he colder northern deserts getting so me species from Arcto - Terti ary sources and the warmer southern d eserts from Ma dro - Tertiary groups. In southe rn Califo rnia, chapa rral vegetatio n, adap ted to t he new Med iterrane an clima te of coo l winters (with so me preci p itation) and long hot summers, dominated the lower mountain areas. During the P leistocene, the cool gla ciation perio ds that left t heir marks o n the Califo rnia mountains allowed temperate vegetation to advance further southward tha n at any other period of the Cenozoic Period. The Redwood Forest moved as far south as Carp enteria and Santa Cruz Island, app roxima tely 200 miles so uth of its pr esent limits. Glaciation during thi s time not onl y produced a cooler climate, but probably a wetter climate. We can see support for this theory from the evidence that many lakes exist e d at that time. Additionally, older pre sent lakes once had greater depths, and even the Mojave Desert shows evidence of many riverbeds that are no longer filled with water. Mono Lake, for example, had obtained a depth of nearly 900 feet during the Pleist ocene Epoch. Owens Lake was 250 fe et higher and overflowed into the basins of China an d Searles lak es, and w ater from the S earles Bas in overflow ed in to the Panamin t Basin t o form a lak e 930 feet deep, which in turn apparently spilled southward into Death Valley, producing a lake 90 miles long and 600 feet deep! Southward migrations of plants during such c older and wetter periods, with subsequent periodic northward migrations, undoubtedly had a great influence on both the evolution and distribution of our present vegetation and the butterflies associated with i t. Much hybridization along the flowering plants would have occurred as species w ere brought into contact repeatedly after long periods of isolation. Movement of species would have occurred up and down the Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 65 mountains, as well as southward and northward. The persistence of over 300 species of Sierran plants in the pine belt of the southern California mountains can best be explained by the extensive southward migrations of plants during the glacial period (Munz and Keck, 1963). The lack of many widely distributed alpine plants in California, found as far sout h as Colorado or New Mexico in the Rocky Mountains but not in the Sierra Nevada, was probably a result of the historical influence of summer dryness in California. On the other hand, the evolution of high - alti tude species fro m somewhat lower - elev ation genera has been notable, with our alpine flora having many endemics. Furthermo re, and pe rhaps mos t signi ficant for the studen t of butterflies, the wh ole regio n has been so broken up into separate mountain r anges and low - lying valleys that a host of ecological niches, often quite isolated from each other today, has been created. Such conditions cannot help but favor endemis m and the develop ment of many ex ceedingly local species and su bspecies of both plants and insects. Thus California has a flo ra whic h is extremely rich in numb er of forms and has many very large genera. The existence of over 7,00 0 species of plants here, and nearly 300 species of butterflies with over 700 subspecies, has its roots in the geological hi story, often orogenic ally violent an d climatic ally dra matic, that w e have traced in th is brief cha pter. B oth plants an d animals in Califo rnia have followe d a diverse series o f pathway s in thei r evolution, but we can conclude with Munz and Keck (1963) that "all have undergone rather rapid and highly polymorph ic differen tiatio n." Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 66 Chapter 3 THE RELAT ION OF THE C ALIFORNI A CLIMATE T O THE BUTT ERFLY FAUN A Because of the influence of its coastal position and mountain relief with a wide range of topography in Califo rnia, th e state inc orporates s everal qu ite differen t climates , with m arked differenc es occurr ing within a very fe w miles. Summe r temp eratures in the low- elevation southeas tern latitud e is not th e chief de terminant of tempera ture but ra ther the iso therms w hich tend to have a north- south tre nd in this state rathe r than the u sual east -wes t trend, as they follow topographic contours of the coastal and Sierran mountains. Most of the United States is marked by sharp differences between summer and winter climates. The midcontinent is flooded by a huge mass of frigid, dry Canadian air in winter, which then moves eastward, often with snow. Winter climate on the West Coast i s quite differen t. A great mas s of relat ively warme r, moist ai r moves south from t he Aleutian Islands o f Alaska, occasionally reaching southern California, which receives nearly all its rain during the winter. Th is Pacif ic air brin gs mild te mperature s alo ng the en tire coast, an d the rain storms are separated by long periods of relatively warm, sunny weather in these winter months. During the s ummer, the Pacific w eather reg ime also dif fers fro m the humid , hot clima te of the eastern states, with their frequent convectional and frontal summer thunderstorms. The moist Aleuti an air masses retrea t northwards, and are r eplaced by mild s table air from the cool subtropical Pacif ic. Virtually no rai n falls on the Pacific coa st in the summer, ex cept in the mountain s. In the in termontan e desert a reas, te mperature s rise to sco rching leve ls. The irr egular topography of Califor nia shatters any general bi oclimatic p attern into an intr icate mosa ic of small clima tic reg ions that di ffer drastic ally accor ding to e levation, exp osure d irection , and sharp variation in soils and vegeta tion cov er. The foll owing brief summary of California' s complex clim ate is based principally on Sprague (1941). The Effect of Temp erature Along the i mmediate coast, th e climate is cool a nd without extreme flu ctuation in temperatu re. This ma ritime - type climate occurs on the lower seaward slop es of the Outer Coast Ranges in the beach du nes of California, w here mild winters and cool summers ar e t he rule. The weather seldom inc ludes freezing tem peratures, nor does f rost occur on the immedia te coast. In fact, on sunny days in January and February, the south-facing slopes and protected sides of the dunes may get quite warm (Langston, 1974). Bec ause of the rela tively warm nights, the development of L epidoptera and oth er insects can start qu ite early in most years. On the other hand, in the summer the coastal slopes and dunes may receive long periods of cold winds and fogs, with the sun not shining for several days to several weeks in a row. During these cool periods, new adult butterflies do not emerge from their pupae and those already in the adult s tage may re main quiescen t, not car rying on their activ ities of nec taring, Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 67 mating, or ovipositing (Langston, 1974). When the weather becomes clear and sunny again for a short tim e, the adult s resume the ir activ ities befo re the nex t siege of su mmer fog. On the eastern slop es of the Coast Ra nges and in the C entral Valley occurs a co ntinental climate, with a greater range in tem perature. Summe r temperatures may exceed 100o F, while winter temp eratures w ill drop below free zing, wit h snow in north ern coun ti es of the Valley. This Valley - lo calized co ntinental c limate give s way to the mountain climate in the Sierra Nevada and other ranges. Langston (1974) summarizes the Inner Coast Ranges and Sierra Nevada climate as b eing characterize d by cold winters and ho t summers, with hi gh te mperatures regularly reaching in the 90's (F). The flight season tends to be shorter in length, particularly at the higher elevations. Below- freezin g temperatu res and w inter frost ten d to be c ommon in the Inner Coast Ranges, while heavy snows and several months of below- freezing temperatures are characteristi c of the Sierra Nevada. Thus the eastern and wes tern slopes of the Coas t Ranges and the Sie rra Nevada as a whole rise to sufficie ntly hig h elevatio ns for sno w; in fact, f reezing temperatu res may o ccur in any month of the year. In areas near the coast, there may be 365 days that are frost - fr ee each year, but above 6,000' in California, fewer than 100 frost-free days occur. In the spring, the days may warm up suddenly, even with night temperatures still remaining below freezing. When the days have warmed, it ge nerally stays w arm and gets progressi vely hotter as summer ap proaches. As a result, a rath er syncronous and n ear total emergence o f adults of a part icular species will occur. With adequate temperatures occurring daily, the adults nectar, mate and oviposit in a short tim e frame. With adult activities concluded, the flight period is soon over in such species (Langston, 1974). The Impact of Precip itation The amount of precipitation in the state is strongly influenced by the distance from the ocean. It is also affected notably by the elevation, shape, and steepness of the mountain slopes, and their direction of orientation in relation to the moisture- b ear ing wind. The prevailing wind from the w est moderat es the cl imate thro ughout t he state and increases p recipi tation. Most of the state is characte rized by an annual w et season and dry season alternation, the w et season falling usually in the cooler months of the w inter. In gen eral, prec ipitation increases f rom south to the north, and is heavier on the southern and western slopes. The average annual precipitation throughout the state varies from less than 2 inches (50 mm) in Death Valley to over 109 inche s in parts of Del N orte County in the nort hwest. However, som e interior desert are as may see entire years pass without moisture, while a recording station in Del Norte County has received as much as 153.54 inches (3,900 mm) and the village of Honeydew, in the redwoods of Humboldt County, has received as much as 174 inches in a year. Mountain recording stations so metimes receive as much as 10 inches (254 mm) in 24 hours, particularly in southern California, with torrenti al floods ensuing. S nowfall may ran ge from a trace o r none near the southern coast to as much as 449 inches (11,405 mm) in the Sierra Nevada. However, great variability in annual precipitation is the rule in California and long drought periods may occur. Occasional wet years, like 1958, 1966, or 1973, may result in remarkably abundant vegetation, where virtually none Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 68 has appeared for years, and great numbers of butterflies, including a number of new records as butterflies fly north from Mexico or Arizona. Cold springs may delay the season as long as three or four weeks in the Sierra Nevada or even in relatively low el evation areas. An import ant clima tic factor fo r both v egetation a nd butterfl ies along th e coast is fog. Summer fog s genera lly increas e with la titude and altitude. Su mmer fogs a re more f requent than in winter along the coast and on windward slopes. However, in winter, fog may be a very important m oisture- de livering sy stem fo r plants in interior areas as in th e Central V alley. HISTO RICAL CLI MATES OF CALI FORNIA The distribution of vegetation and butterflies in California has been anything but static in recent his tory. D ry periods alt ernatin g with wet p eriods were tied to the advance an d withdraw al of the great Pleistocene glaciers during the 100,000 years. Thes e vegetational ch anges have occurred down to rela tively re cent time s. For examp le, by st udying the f ossil feca l material in coporolite deposits of the extinct ground sloth, Nothrotherium shastense , and of the remarkably ubiquitous wood -rat middens deposits, containing abundant well- preserved plant macrofossils, Wells and Berger (1967) showed that at the close of the Wisconsin glacial period, there was widespread expansion of the pinyon-juniper zone downward into areas that are presently desert. This paleo botanical record, conveniently kept by sloths and wood-rat middens in protected caves, show that xerophytic juniper -woodlands vegetation descended to an elevation of 1,100 meters around 9,000-10,000 years ago, which is some 600 meters below the present low er limit of woodlan d (1,700 me ters) in the lati tude of Fren chman Flat i n the Mojav e Desert. D esert or semi - desert shrubs co -existed with the woodland trees throughout much of the elevational span corresponding to this weather-lowering of the pinyon-juniper woodland. Joshua Trees, accompanied by desert shrubs, moved down as low as 600 meters at Gypsum Cave, Nevada. These discoveries o f macrofossil evi dence for movement s of the trees dow nward during wet periods in Californ ian histo ry hav e important implica tions for but terfly dis tributio n patterns. For example, they show that there was a former continuity of the many disjunct stands of vegetation in the Mojave Desert region, at least along the higher divides connecting most of the ranges. This suggests that some species, su ch as Papilio indra which today exists in highly isolated populations in the higher parts of the Mojave Desert, may have had a more continuous range as recently as 9,000 years ago. Subsequent isolation by the juniper- dominated woodland shifting u pward on lime stone moun tains, as th e clima te of the are a became h istorically drier, le ft small, separate d popul ations behind of both larval foodplants and butterflies. In these butterfly colonies, gen etic drift and founder e ffects could operat e with relative ease to produce dramatic evolutionary changes in phenotype, resulting in such different subspecies a s the narro w -banded P. indra martini and the wide-banded P. i. fordi being located in mountain ranges only twenty miles apart today. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 69 Chapter 4 PLANT COMM UNIT IES OF CALIFO RNIA The plant community concept is a somewhat artificial one because of t he continual change in phases of any one community, depending upon w hich species is or are dominant. Other problems in clude the transit ional ecotone area s between co mmunities, etc. Phi llip A. Munz (1959), in his book A CALIFORNIA FLORA , presented an excell ent scheme, bu t other authors have identified different divisions, and in fact, there are probably an almost infinite variety of communities that cou ld be defined in a state as geograp hicall y large and as topogr aphically diverse as California. Because of its widespread adoption, utility, and r eady recognition, the Munz scheme of plant communities has been adopted in the present book to identify the basic plant communities in which the butterfl ies of Cali fornia are fo und. In each of the fol lowing plant communities, we follow an outli ne of naming the plant community , then listin g the domi nant plants that charac terize that commun ity, the butterfl y foodplan ts that are fo und in th at communi ty, and th e charact eristic b utterfli e s that usually occu r in that plant community. A brief discussion follows in each community section. 1. COASTA L DUNE SAND PLA NT DOMIN ANT OR CHARA CTERIS TIC PL ANTS Ammophil a arenaria, Bromus maritim us, Elymus spp., Poado uglasi i, Junc us les ueurii and Car ex pans a; species of Abr onia, C arpobr otus, Mesembry anthe - mum, and Cakile;Calys tegia soldanella, Eschs cholzia californica mariti ma, Fr agari a chil oensi s, Ambr osi a chami ssonis, Artemi sia py cnoce phala, Atr iplex l euco phyll a, Cami ssoni a cheiranth ifolia, Lathyrus litto ralis, and shr ubby spec ies of Lupinus . BUTTER FLY FOODP LANTS Castilleja mollis, Core throgyne filaginifolia , Eriogonum latifolium, E. parvifo lium, Lotus scoparius (prostrate ecotype), Lupin us arbor eus, L. ch amissoni s, Viol a adunc a . CHARACTERIS TIC BUTTERFLIES Thessalia leanira ssp. (Oso Fl aco dunes ), Chl osyne g abbii , Speyeria zerene myr tleae, S. z. near hippolyta , Apodemia mormo virgulti ( El Segun do dunes e cotype), A . mormo ssp. (Mari na dune s, Mo ntere y Co.), Call ophrys dumet orum, C . perplexa perplexa, Glaucopsyche lygdamus near australis (Mari na dunes ), G. xe rces, Euphil otes bat toide s allyni , E. enoptes near tildeni (Marina dunes). DI SCUSSION This pl ant c ommu nity i s co nfined to sa ndy beac hes and sa nd dunes scattered along the coast of Cali fornia. Annual preci pitat ion range s from m oderat e to hea vy, mostl y durin g the winter mont hs, b ut co nside rabl e fog during the s umme r mon ths tends t o suppl eme nt this moisture. Temperature fluctuation s tend to be mi nimal , both se asona lly and diurn ally. T he gro wing season is 12 months long, and freezes are extremely rare or non - existent. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 70 The but terfl y inhabi tants o f this commu nity t end to have long flight periods b ecause of the mild te mperat ures and m oderat ing influe nce of t he oce an; most are s pring o r earl y summe r flyi ng (see Lan gston, 19 ). Becau se of the extremely restricted geographic area of this particular p lant community, most of the endemic butterfly subspecies inhabiting it ar e restricted in th eir distribution , and several have been declared endangered species as a result. In fact, o ne butt erfly specie s and two s ubspeci es wit hin t his pl ant commu nity ha ve becom e exti nct as t he resul t of h uman encroa chme nt upon i t: Cercyonis sthenele sthenele, Glaucopsyche xerces , and Icarici a icari oid es phere s . Lar g e areas of this habitat have be e n d estroyed by urbaniza tion and other human a ctivi ties . In parti cular, t he cit y of Sa n Franc isco has re place d an ext remel y ext ensive c oasta l dune habit at, a nd the western coastal area of greater Los Angeles has similarly destr oyed o ver 95% of the once extremel y ext ensi ve sand dune system s in tha t geogra phic area. Ad ditio nally, ot her e xtensi ve sand d une habi tats , such as th e Oso Fla co Sa nd Dunes in San Luis O bispo C ounty and the Ma rina Dunes in M ontere y Count y, have been ext ensive ly alt ered by such h uman a ctivi ties as off - road vehicle activity , military reserv ations, sand - mini ng, and enc roachm ent o f exoti c specie s of pla nts s uch as Ic e Pl ant ( Mese mbrea nthum species). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 77 7. NORTHERN CO ASTA L SCRUB DOMIN ANT OR CHARA CTERIS TIC PL ANTS Artemisia californica , Baccharis pilu laris , Cean othus thrysifloru s repens , Dipla cus aura ntiacus , Eriogonu m parvifolium , Eriophy llum staechadifolium , Gau ltheria shallon , Rubus spp., Achillea bo realis californ ica , Amsinckia spectabilis , Anaphali s margari tace a , An gelica he nders onii , Castilleja latifo lia , E riger on glauc us , Grindelia stricta venul osa , Hera cleum lana tum , Iris do uglasiana , Lup inus spp., Sisyrinchium bellum , Armeria maritima californica , Dudleya farinosa , Plantago maritima californica , and Spergularia macrotheca . BUTTERFLY FOODPLANTS As ter chilensis , Ana phalis ma rgari tacea , Angelica hendersonii , Aster chilensis , Castilleja spp., Ceanothus thyrsiflorus , Diplac us aur anti acus , Eriogonum latifoliu m , E. parvifolium , Heracleum maximum , Lupinus a lbifrons s sp. collinus , Scrop hularia cal ifornic a , Sedum spathulifolium . CHARACTERIS TIC BUTTERFLIES Papilio zelicaon , Euphy dryas chalce dona c halce dona , E. editha baro ni , Phyciodes c ampestris campestris , I ncisal ia mossi i bayens is , I. m. doudorof fi , Callop hrys dumetor um , Lycaena heteronea heteronea , Euph ilotes enoptes smithi , Icaricia icarioides m issionensis . DISCUSSION This pl ant c ommuni ty occ upies a rathe r narr ow coas tal s trip from San Lu is Obispo County nor th to the Oregon bord er, where i t is l ocate d above the Coa stal Dune S and pl ant commu nity bu t below t he re dwood f orest and othe r coa stal forest communities. It is la rgely below 500 feet in elev ation, and a verage rai nfall r anges from 25 to 75 inc hes, with consi derable fog an d wind. There is a lo ng gro wing se ason , from 10 t o 12 m onths, a nd te mperat ure fl uctuati ons bot h diurnal ly a nd se asonal ly te nd to be grea tly m odera ted by the proximity to the oce an. The plants in this community tend to be low - growing bu t often forming a fairly de nse vegetation cover, sometimes with small areas of grassland within it. Because of the frequent location of this pl ant c ommu nity o n stee p slope s near the oc ean, t here ha s b een somewhat less impact on it by human activities. However, one of the most ext ensi ve area s of t his pl ant c ommuni ty, o n San Bruno Mounta in in Sa n Ma teo Co unty, is threat ened with encroa chin g urbani zatio n fro m the Sa n Fra ncisco metrop olita n area. At least 5 butter fly species have d eveloped distinctive subspecies confined to this plan t community: two of these are restricted enough to be classified as endangered subspecies. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 78 8. SOUTHERN CO ASTA L SAGE SCRUB DO MIN ANT OR CHARA CTERIS TIC PL ANTS Artemisia californica , Crossoso ma cal iforni cum , Di placus , Encelia californ ica , Eriogo num fas cicul atum f ascicul atum , Galium , Haplopappus , Helianthemum , Lotus , L upin us , Malacotham nus , Opuntia , Mal osma la urina , Rhus integrifolia , Salvia , Sen ecio , Solanum , Toxi codendr on r adicans diversilobum , Marah macr ocarpus , Cal ystegi a macrost egia , and species of Castilleja , Haplop appus , Lot us , Lupinus , and Zauschn eria , and Er iophyllum confertiflor um . BU TTERFLY FOODPLANTS Astragal us tricho podus ssp. leucopsis , Caulant hus lasiophyllu s , Desc urainea pinnat a , Dipl acus spp., Encelia californ ica , Erio gonum c inereum , E. fasciculatum , Lotus argophy llus , L. scopa rius . C HARACTERISTIC BUTTERF LIES Anth ocharis cethur a cat alina , A. sara gunde ri , Euphydryas chalce dona c halc edona , Char idryas gabbii , A podemia mor mo virgulti , Calephelis n emesis californica , Glauc opsyc he lygdam us aust ralis , G. l. pal osve rdesens is , Euphil otes batt oide s bernar dino ( Er iogon um cine reum ecot ype). D ISCUSSIO N Under this plant c ommu nity, we lum p 3 o f the s ubdi visions of Farn (19 76): Sea - bluff Succ ulent , Marit ime Sage Scrub, and Maritime Desert Scrub . This plant community is found in the southern third of the state on the islands and the immediate coast u p to t he chaparral belt of th e coastal moun tains. It is fairly r ich in plant specie s beca use of the favo rable c onditi ons of a long growing season, moderate tem peratures, moderate winter rainfal l, and t he mo derat ing effe cts of s ummer f og. Most of the associated butterfly species ten d to be spring or early summer flier s, in keeping with th e period of time at wh ich the plants in thi s c ommuni ty are acti vely gr owing a nd fl oweri ng. A s is the case with the other coastal Californ ia communities, large tracts of this habitat along the coast have been replaced by housi ng devel opments . On the Cali fornia Is lands, the re has been c onside rable degrada tion o f this ha bita t from gr azing b y fera l animals, although programs are now in place to eliminate this source of habitat d egradation on all the Californ ia Islands. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 79 9. INLAND SA GE SCRUB DO MIN ANT OR CHARA CTERIS TIC PL ANTS Artemisia californica , Dipl acus spp., Ence lia fari nosa , Eriog onum fasc iculat um , Eriophyllum confer tiflorum , Haplopa pppus spp. , Helianthenum sc oparium , Eriodi cty on trichocalyx , Lotus s copari us , Lupinu s and Malac othamn us spp., Malos ma lauri na , Opuntia littora lis and O. parryi , Rhus , Rhamnus , and Salvia spp., Solanum xanti , and Toxi coden dron radic ans div ersil obum . In sandy wash es, Cucurbita foetidissim a , Lepidos partum squamat um , Rham nus spp ., and many a nnuals . BU TTERFLY FOODPLANTS: Be bbia juncea , Caul anthus simul ans , C orethr ogyne filaginifo lia , Dudleya lan ceolata , Eriog onum elonga tum , E. fasci culat um , Lotus scop ariu s , Plantago spp. C HARACTERISTIC BUTTERF LIES Anth ocharis cethur a ceth ura , Euphy dryas edith a qui no , Chlosy ne gab bii , Apodemia mor mo vir gulti (inla nd ec otype) , Calephelis wrig hti , Callophrys perplexa perplexa , L ycae na gorgon go rgon , Glauc opsyc he lygda mus aus trali s , Philotes sonorensis , Eup hilotes batto ides bern ardino , Hesperia columbi a . D ISCUSSIO N This community is closely related to the preceding one, in having a similar species composition , although it tends to b e subject to much greater extremes of temperature and lower precipitation as a consequence of its distance from the coast. This c ommu nity is usuall y pre sent on s andy , grave lly, o r roc ky slopes on t he coa stal s ide of the mou ntains from t he so uth Coast Ranges in San Luis Obispo Coun ty to Baja California. Elevation ranges from near sea level to approximately 3000 feet, and annual precipitati on i s 10 t o 20 i nches , with a gro wing seaso n of 8 to 12 m onths. Wi nter mi nimum tempe ratu res ma y reach freezi ng, al thou gh onl y for brief periods of t ime, while summer maximum temperatures are often above 90 o F. T he dom inant pla nts in this c ommu nity te nd t o be s mall shrubs and for m a mo re o pen com munit y than t hat s een in the c haparra l communities. Most of the butterflies of this particular commu nity fl y duri ng t he sp ring m onths when t he pl ants are eithe r in bl oo m or puttin g out ne w growth. By summe r, mos t of the plant species are in a dormant state, and likewise there are ve ry few butte rfly spe cies flying during t his s eason or in t he fall months. A ltho ugh muc h of t he rea l es tate de velop ment c urrent ly ta king place in southern Californ ia occurs within this plant community outsi de of the vall ey bottom s, it occupie s a rathe r exten sive geographic area and is not immediately t hreatened. However, in nor thern San Diego County, w estern Riverside County, and southw ester n San B ernardi no County , real e sta te devel opme nt is accelerating at such a pace that within the next few years significant d estruction or even virtual elimination o f this plant commu nity i n that area could take place. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 80 10 COASTA L CLOSED - CONE CO NIFERO US WOODLAND DO MIN ANT OR CHARA CTERIS TIC PL ANTS P inus m uricat a and its variety remorata , P. radi ata , Cupress us macroc arpa , C. abr amsi ana , C. goveniana , and C. guadal upensi s . BU TTERFLY FOODPLANTS Pinus muri cata , Horkelia sp. C HARACTERISTIC BUTTERF LIES Incis alia e rypho ne near shelton ensis , Pyrgus ruralis ruralis (Mont erey Pe ninsul a). D ISCUSSIO N This was pr obabl y once an extensi ve com munit y wit hin t he state, but is currently confin ed to only a few scattered locations along t he coa st an d on the Cal ifornia Isla nd. It is dom inate d by closed - cone coni fers, a nd freq uentl y has an un derstor y of scleroph yllous chapa rral shrubs. In Mendo cino C ounty a few miles inland from the Pacific coast, there is a dwarfed phase of this co mmunit y formi ng a rat her dist incti ve pigm y forest , where trees can bear cones and are only one or two feet tall, as a result of the e xtrem ely po or soil s on whi ch the y grow . There are no butter fly spe cies unique to th is comm unit y but a s mall numbe r seem at least loosely as sociated with it. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 81 11. INLA ND CLOSED - CONE C ONIFERO US WOODLA ND DOM IN ANT OR CHARA CTERIS TIC PL ANTS Pinus attenuata , Cupress us mac nabian a , C. sargentii , C. arizonica , C. bakeri , C. guada lupensis . BUTTER FLY FOODP LANTS Cupres sus g uadalu pensi s ssp . f orbe sii , C. sargentii , C. macnabi ana ( ?), C. nevadens is (?), C. bakeri (?). Mitour a barryi barryi , M. muiri , M. thor nei . DI SCUSSION This is the inl and re prese ntati on of the Close d - cone Conife rous Forest , and is usual ly fou nd on eit her se rpent ine or volcani c soils , and i s loose ly ass ociat ed wit h chapa rral , oak wo odla nd, or eve n a sp arse yel low pine f orest. It is f ound i n seve ral wi dely scattered locatio ns throughout the state, with the largest s tands of thes e uni que co nifers i n the North C oast R anges and ext reme northe rn Sie rra Ne vada and M odoc Pl ateau regio ns. Un ique to these stands of conifers are several species of Mitoura . The coni fer s pecie s are al l pyrop hytic , requi ring regula r fires for ope ning of t heir co nes to rel ease see ds; the effect of these recurrent fires on the Mitoura popula tions is unstu died at this t ime, b ut may s hed so me light on the curr ent distributio n patterns of these species. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 82 12. SHORE PI NE WOODLA ND DOMIN ANT OR CHARA CTERIS TIC PL ANTS P inus c ontort a co ntorta , Arctos taphyl os uva -ursi , G arrya elliptica , Gaultheria shallo n , Myrica californica , and Vaccini um ov atum . BUTTERFLY FOODPLANTS Pin us contorta , Arctostaphylos uva -ursi . CHARACTERIS TIC BUTTERFLIES Incis alia ery phon ne ar sheltonens is , I. a ugust us iroi des . DISCUSSION This is a rather limited community within the state, being found only at severa l loc ations along the n orther n coast from near Cape Me ndoci no nort hward into Or egon. It is usua lly fo und on stabi lized s and du nes or c oasta l bluff s, and consis ts of a relativel y low open woo dland domi nate d by Pinus contorta contor ta . Within the state, perhaps th e best development of this plant c ommu nity is on the Samoa P eni nsula we st of Humbol dt Bay in H umbol dt Cou nty. Onl y two b utterfl y speci es se em associated with it, bo th u nivoltine spring - fliers . Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 83 13. TORREY PI NE WO ODLAND DO MIN ANT OR CHARA CTERIS TIC PL ANTS Pi nus torreyana . BU TTERFLY FOODPLANTS Undetermi ned. C HARACTERISTIC BUTTERF LIES Undet ermi ned. D ISCUSSIO N This is a coas tal woodland community pr esent in only two very limi ted sta nds, one in Sa n Die go Cou nty nea r Del Ma r an d the other o n Sant a Rosa Isl and. Thi s comm unity ha s not been studie d from t he sta ndpoint of ass ociate d butt erfly s pecies and bears invest igation. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 84 14. NORTH CO ASTAL CO NIFEROUS FO REST DOMIN ANT OR CHARA CTERIS TIC PL ANTS A bies gran dis , Chamae cypar is laws onian a , Pic ea sitchensis , Pseudotsuga menziesi i , Thu ja plicata , an d Tsuga he tero phyll a . The de nse un derst ory incl udes Acer circinatum , Rhamnus purshi ana , Poly stichu m munit um , an d many ot her spe cies with Selagi nella or egan a often e piphyti c on t he moss - hung tree trunks and b ranche s (Th orne, 1976). BUTTERFLY FOODPLANTS Calamagr ostis p urpuras cens , Cardami ne cali fornic a , Ca re x sp. , Pseudotsuga me nziesii . CHARACTERIS TIC BUTTERFLIES Neophas ia menapia melan ica , Pieris napi marginalis , Oen ei s nevade nsis i duna , Euphyes vest ris osceola , Carteroce phal us palaem on ro bustus . DISCUSSION This is a coastal co niferous forest of northern Califo rnia ranging from scatter ed localities in Sono ma County northw ard to the Oreg on borde r, fr om sea l evel to appr oxima tely 1 ,000 fe et eleva tion o n the out ermos t Klam ath and C oas t Ranges . It i s a very de ns e, tall, wet fo rest dominated by conifers o ften reaching 150 t o 200 fe et in he ight. The re is al so a dense underst ory of shrubs. It is f ound in th e area of highest rainf all in the state, with some station s receiving 110 inches of precipitation a y ear, with f reque nt summe r fog. It has a lon g growi ng seas on, wi th cool a nd equa ble temperat ures . The de nser portio ns of this fores t are proba bly devoi d of butter fly s pecies , and m ost of the speci es rec orde d for this particular p lant community are found in openings with in the forest o r along roads, o r along wat ercou rses where the for est canop y is abs e nt. Because of marginal weat her conditions for obser ving butt erfli es in this commu nity for much of the y ear, little research has b een done on it. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 85 15. REDWO OD FOREST DOM IN ANT OR CHARA CTERIS TIC PL ANTS Sequoia sempervirens , Pseud otsug a menzi esii , Torreya californ ica , Acer mac rophy llum , A lnus or egana , Arbu tu s menziesii , Quercus densiflora , Umbellularia ca lifornica , shrubby s pecies of Rhod odendron and Vacc inium , C eanot hus thyrsifloru s , Coryl us cornut a calif ornic a , Euonym us occidentalis , Gault heria s hall on , Myrica californica , Ro sa gymnocar pa , and species of Rubus , herbs suc h as Asarum caudatum , Oxalis o regana , Viola sempervirens , and species of the Berbe ridace ae , Ranu nculac eae , Liliaceae , and Orchidaceae , and fer ns like Adiantum pedat um aleuti cum , Athyrium filix - femina californ icum , Blechnum s picant , Polystichum munitum , an d Woodwar dia f imbriat a (Thorn e, 1976). BUTTER FLY FOODP LANTS Card amine angul ata , C. californ ica , Dicentra formo sa , Rib es divaricatum , Sedum spathulifolium , Viola se mperv irens . CH ARACTERIS TIC BUTTERFLIES Parnas sius c lodi us str ohbe eni , Pieris napi sequoia , P. n. venosa , Boloria ep ithore epithore , Polyg onia f aunus fulvescens , P. oreas , Incisalia mos sii marinensis , Celastria argiol us ssp. (N. Coast segrega te), Euphyes vestris near osceol a (Santa Cru z Mts.). DI SCUSSION This is anot her hi gh, de nse, c oastal fores t of t he nort hern half of the state. It is found from southern Mon terey County northw ard to southern Or egon. It is found from ne ar sea level to as hi gh as 30 00 feet on the seawar d slope of the o uter Klamath and C oast Ra nges, a nd re ceive s hea vy wint er rai n an d freque nt den se, d rippin g fog during the s ummer months . In de nser port ions of this fore st, but terfli es are virtua lly a bsent, but in clearings and along watercourses, trails, or roads, butter flies may be freque ntly e ncounte red. T he spec ies compo sition of thi s plant com munity i s best kno wn thro ugh the field wor k of both J. W. Tilden a nd O. Elt on Sette , resident lepido pteris ts of t he San F ranci sco Ba y regio n who m ade freque nt col lect ing tri ps into t he r edwoo d fore sts of t he s outher n Santa Cruz Mo untains. At leas t one subspecies u nique to this habitat ( Pa rna ssiu s clodius strobin i ) has poss ibly bec ome extinct due to destru ction of its habitat in the Santa Cruz Mount ains. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 86 16. DOUGLA S FIR FOREST DOMIN ANT OR CHARA CTERIS TIC PL ANTS Dominated by Pseudotsuga menziesii ; often associated with the Douglas - fir a re Quer cus de nsi flora , Chrysol epis chrys ophyll a , Arbutus menziesii , Ceanot hus t hyrs iflor us , Dipl acus aurant iacus , Lonicer a hispi dula vacill ans , Rhamnus californ ica , and Toxic odend ron ra dicans div ersilob um . BUTTERFLY FOODPLANTS Ca rdamine californica , Ceanothus thyrsiflorus , Pseudotsuga menziesii , Quercus de nsifl ora , Rhamnus californica , Sedum spathulifoliu m . CHARACTERIS TIC BUTTERFLIES Papil io eury medon , Neophas ia men apia me lanica , Pieris napi marginalis , P. n. venosa , Inc isal ia mossi i marine nsis . DISCUSSION This is a fo rest d omina ted by Dougl as Fi r, foun d in t he N orth Coast Ranges and Klam ath Mo untains Asse mblage gener ally inland fro m the redwood fores t. It is found up to 4500 feet in elevation, and sometimes forms nearly pure stands of Douglas Fir 200 feet in height. There is an understory of smaller trees and sh rubs, b ut wit h less di vers ity than t hat o f the o ften a djacent Northe rn Mi xed Eve rgree n fore st. T his fo rest has bee n po orly studied, and what little data we h ave is the result of field s tudies prima rily i n smal l patche s of t his f orest i n Mari n and S onoma counties in the Nor th Coast Ranges. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 93 23. FOOTHILL WOODLAND DOMIN ANT OR CHARA CTERIS TIC PL ANTS " Upslope fro m the g rassla nd it is a pa rkli ke oa k woodl and ("sava nna") dominate d by Quer cus l obata , Q. agrifolia , Q. dougl asii , Q. wislizen ii , Q. chrysol epis , and Aesculus californ ica . At hi gher el evat ions the woodl and bec omes de nse r with d omina nce by Querc us dougl asii , Aesculus californica , and Pinus sab iniana , and cons iderabl e unde rbrus h of such shrubs as species of Ceanot hus and Arctost aphyl os , Eriodic tyon californ icum , and Rhamn us californica tomentella . In th e mesic valleys, it grad es into riparian woodland with a rich mixtu re of s hrubs , suc h as Calycanthus occid entalis , Cercis occcidenta lis , Fraxi nus dipe tala , Myrica h artwegii , Philadelphu s lewisii , Ptelea crenula ta , Staphy lea bolanderi , Umbellularia californica , and even such localized endemics as Carpenteria califor nica (Fresno Cou nty)" (T horne, 1976) . BUTTERFLY FOODPLANTS Arabis sp p., Ar ceuthob ium campylopodum , Cardamine californ ica , Ce anothus spp., Collinsia tincto ria , Keckiella breviflora , K. lemmonii , Ptele a crenul ata , Quercus spp., Rhamn us cal ifornic a , Sed um spat hulif olium . CHARACTERIS TIC BUTTERFLIES Papilio eu rymedon , P. multicaudatus , Pontia sisymb rii sisymbrii , Pieris nap i microstriata , Antho charis sara sara , A. lance olata l anceol ata , Coenony mpha cal ifor nia cal iforni a , Cercyonis sthen ele silvestris , Adelpha bredowii californica , Euphy dryas c halce dona chalcedona , E . c. dwinellei , E. edit ha rubic unda , Speyeria callippe rupestris , Habrodai s grunus grunus , Sa tyrium califo rnica californ ica , S. auretorum auret orum , S. s aepium s aepium , Incisalia m ossii windi , Mitour a joh nsoni , Ochlo des sy lvanoi des sy lvanoi des , O. agric ola ne morum . DISCUSSION This c ommuni ty is cl osel y rela ted to t he No rther n and So uther n Oak Woodland communities, and forms a transitional woodl and bet ween t he Grass land a nd Cha parra l comm unitie s at its l ower e dge and t he ye llow Pine Forest and M ontane Chaparral commun ities at its upper e dge. T his is a much m ore florist ical ly ric h commu nity t han the Northe rn or S outher n Oak Woodl and co mmuni ties, a nd in fact ma y be s ubject to furt her subdivision as it is studied in greater detail. It is found on the lower slopes of the inner Coast Ran ges and the eastern slope of the out er Coa st R anges t hrough t he we stern foothi lls of the Sierra Nevada and i nto the northe rnmost part of the Trans verse Ranges . It has a broa d ele vation r ange, from 400 t o 5000 feet elevation. There is a much greater number of butterfly species associated with this plant community, due to its rich flora as well as more divers e habit ats of ca nyon bot toms, rocky s lope s, an d extens ive interface s with adjacent plant communities. As with the Northe rn and Southe rn Oak W oodla nd com munit ies, t he butter fly spe cies of this c ommuni ty are l argel y spring or earl y summer univoltine species. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 94 24. MIXED CHAPARRA L DO MIN ANT OR CHARA CTERIS TIC PL ANTS T he domi nant shrubs a re us ually s pecie s of A denost oma , Arctostaphylo s , Ceanothus , G arry a , Que rcu s , Rhamnus , and Rhus , plus C ercoc arpus b atuloi des , Dendro mecon rigid a , Fremont oden dron c alif ornicum , Heteromeles arbutifolia , Picke ringi a montan a , Prunus ilicifolia ilicifolia , Tricho stema lanatum , and Yucca w hippl ei , but the species co mposition varies greatly with the various sub - comm uniti es (Tho rne, 1976). BU TTERFLY FOODPLANTS Adenostom a fas ciculat um , Ar abis spp., Astraga lus douglasii , Castilleja foliolo sa , Ceanot hus spp., Cercocar pus betul oides , Eriogon um fasci culat um , Fremont oden dron cal ifor nicum , Keckiella terna tus , Ko eleria cristata , Lotus scop arius , Malacotham nus spp ., Prunus ilicifolia , Quercus du mosa , Rhamn us cal ifornic a , R. cr oce a , Tausc hia argut a . C HARACTERISTIC BUTTERF LIES Papilio zelicaon , P. indra per gamus , P. eur ymed on , Anthoc haris lanceol ata aust rali s , C olias harfor dii , Euphydr yas chalced ona chalcedo na , Thessa lia leanira leanira , T. l. wrighti , Satyrium californ ica californica , S. aur etor um auret orum , S. a. spadix , S. saepium s aepiu m , S . s. chalcis , S. s. c hlorop hora , S . s. f ulvescens , S. tetra , Incisal ia augus tus iroides , Lycaena hermes , Celas trina ar giolus echol , Glauc opsyc he ly gdamus australis , Euphi lotes battoi des ber nardin o , E. en optes cry ptorufies , Hesperia colum bia , Ochlo des agric ola agricola , O. a. nemo rum , Erynnis brizo lacustra , E. pacuv ius cal lidus , E. afrani us , Heliopetes ericetorum . D ISCUSSIO N This commun ity of scle rophyl lous shru bs ada pted t o a Mediterranean - type climate, is widespread on the slopes of the foothills and mountains of Californ ia from Shasta Cou nty southward into Baja Califor nia. It has an annual rainfall of 14 to 25 inc hes, vi rt ually all of it falling during the cool winter months , and i t is subject to ho t, dry s ummers . The comm unity is fire - adapted, and many of its species require fire for seed germi natio n and for re juve nation of senesce nt pla nts. Ma ny of the annua ls in this community will only sprou t after burns. T his community h as a relatively r ich butterfly f auna, with many specie s (pa rticul arly the cha parral ha irst reaks) being l arge ly associated with it and to a much lesser extent with other comuni ties. Mo st speci es are univolt ine, wit h peak fl ight activity du ring t he sprin g and early s ummer . Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 95 25. SERPENT INE CHAPA RRAL DOMIN ANT OR CHARA CTERIS TIC PL ANTS Que rcus durata , Gar rya buxifolia , G. congdonii , Ceanothus diverge ns c onfusa , C. ferrisae , C. jepsonii , C. pumilus , Arctos taphy los mo ntana , A. obispoensis , and certain subspe cies of A. can esc ens , A . hookeri , and A. st anfor diana . BUTTERFLY FOODPLANTS Adenostom a fasci culat um , Arceuthobium camp ylopodum , Ceanot hus sp p., Koeleria cr istata , Lomati um margi natum , Pedicul aris densif lora , Querc us durata , Rhamnus californica , Strept anthus spp. CHARACTERIS TIC BUTTERFLIES Papilio zelicaon , P. ind ra s sp. (NW Ca liforn ia segr egate), Pontia sisymbrii sisymbri i , Euchl oe hyant is hy antis , Euphy dryas e ditha l uesthe rae , Satyrium saepi um chal cis , Mitour a muiri , M. john soni , Hesperia c olumbi a , Erynnis brizo lacustra . DISCUSSION This s ubcomm unity of cha parral is f ound o n stone y, in fert ile serpe ntine soils and is readi ly sepa rable from Mixe d Chap arral communities by its c onstellation of certain serp entine endemic plants which are often acc ompanied by more widespread chaparral speci es. G rowi ng wi th thes e cha parral plant s are also sometimes found the serpentine endemic closed - cone cypresses, such as Cupre ssus sarge ntii and C. macnabiana . I t is foun d prima rily i n the no rther n two - t hirds of the sta te, in the Coast Ranges a nd Sier ra Ne vada F oothil ls, at a num ber of widely scatter ed localities. The but terfl y fauna of se rpent ine cha parral habi tats in the North Coast Ranges has been stu died by Har rison a nd Shapir o (19 ). Several butterfly subspecies or ecotypes are virtually completely restricted to serpentine, such as Mitoura muiri or Erynnis brizo lacustr a (at leas t for the nor thern Californ ia popul ations of this skippe r). The C hecke rspot, E uphydryas editha luest herae , is largely conf ined to serpentine habitats, even though its foodplan t may be found on no n - serp en tin e substrates a s well; this suggests th at features of th e micro - habita t cont roll ed by the serpe ntine s ubst rate ma y be as important as the availability of the foodplant itself fo r the prese nce of t his b utterfl y. The butterflies asso ciated with this plant commun ity are largely univol tine, with m ost fli ght pe riods occur ring du ring t he months of April a nd May. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 96 26. MONTA NE CHAPARRA L DOMIN ANT OR CHARA CTERIS TIC PL ANTS Chrysole pis sempervirens , Querc us densi flora ec hinoi des , Q. wislizenii frutes cens , Q. sadleria na , C. chrysol epis nan a , Q . vaccinifo lia , Ceanothus cor dulatus , Arctostaphylos parry ana , A. pat ula , A. punge ns , A. pringlei , and Ch amaebat ia fol iolos a . BUTTERFLY FOODPLANTS Ceanothus cord ulatus , Que rcus c hrysol epis n ana , Q. vaccinifo lia , Q. wislizenii frutescens . CHARACTERIS TIC BUTTERFLIES Papil io eur ymedon , Ery nnis propertius , E. pacuvius liliu s . DISCUSSION This is a relativ ely high - elevation chaparr al community which is us ually found wi thin l arge o penin gs of t he mon tane coniferous forest. The occupation of these open areas by a chapa rral com muni ty may b e the res ult of recurre nt fire s, or possi bly eda phic fa cto rs which pre vent coloni zation by fores t trees. Relative to the lower elevation chap arral communities, this o ne is s omewha t depa uperat e in s pecie s, and o ften s tands of it wi ll be c ompos ed pri maril y of one o r two s pecie s of shr ub. Likewise, the butterfly fauna is not particular ly remarkable, consi sting of speci es whic h are al so fou nd in the adjac ent coniferous fore sts. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 97 27. DESERT TRANSITI ON CHAPARRA L DOMIN ANT OR CHARA CTERIS TIC PL ANTS Que rcus corn elius - melleri , Quercus turbi nell a , Ceanot hus greggi i , Arctosta phylo s pungens , P urshia glandul osa , and Purshi a sta nsburi ana : often accompanie d by Arctost aphyl os glauca , Ceanot hus leuc oder mis , C. crassifolius , C. cune atus , Garrya fla vescens pallida , Fremontode ndr on cali fornic um , Cercocarpus betuloides , an d Yucca w hippl ei . BUTTERFLY FOODPLANTS Arabis perennans , Ce anot hus greg gii , C. cune atus , Cercocarpus betuloides , Dale a searl siae , Descur ainea pi nnat a , Dudleya ab ramsii , Eriog onum elonga tum , E. f ascicul atum , E. wrightii , Frem ontode ndron californicum , Purshi a glan dulosa , P. stansbu riana , Querc us cor nelius - melleri , Q. turbi nella . CHARACTERIS TIC BUTTERFLIES Euchl oe hyant is sp p. (Peni nsula r Ra nges se grega te), Anthoc haris l anceol ata dese rtolimb us , Zerene cesonia , Euphy dryas c halce dona c orrale nsis , E. c. h enei , Satyrium saepi um fulv escens , S . t etra , S. behrii , Incisalia fotis mojavensis , Ph ilotes sonorensis , Euphilotes battoides bernar dino , E. enoptes dammersi , Heliopetes ericetorum . DISCUSSION This is a chapa rral c ommuni ty of t he tra nsmont ane sl opes o f the mounta ins, from the s outhe rn Sierr a Neva da alo ng the n orth slope of the T ransve rse Ra nges, a nd sout h alon g the eas tern slopes of the P enins ular Ra nges. It is als o foun d in th e highe r mountain s of Jos hua Tre e Nat ional Monume nt an d in t he highe r parts of the eas tern Mojave Desert. Its p lant species composition is similar to that of th e Cismontan e Mixed Chapa rral co mmuni ty, but some of the domi nant specie s diffe r as well as the relat ive percentages of the various species composing it. The butterfly species composition of this community is similar to that of mixed chaparral but with several distin ctive subspec ies whi ch sh ow the de sert in fluenc e on t his com munit y. Because of its strategic location between the mountains and the desert, m any species are taken in this community which are actually strays from adjacent hab itats. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 98 28. SERPENT INE DWA RF PINE WOO DLAND DOMIN ANT OR CHARA CTERIS TIC PL ANTS Pinus monticola , Junipe rus c ommunis saxat ilis , Lithocarpus densiflora echn ioides , Arctostaphylo s spp., Ce anothus sp p., Castanops is , Rham nus , Hapl opappus lance olatus in th e drier portion s; in moist to boggy areas , Darlingtonia , P hlox , Orthoc arpus , Mimul us , Ranunculus , Vicia , and Vi ola spp. BUTTERFLY FOODPLANTS A rceuthob ium campylopod um , Castilleja brevilobata , Ceanot hus sp ., Collinsia linear is , E riogonum ternatum , Rhamn us cal ifornic a , Se dum lax um perple xum , Sidalcea s p., Strept anthus h owell ii , Tausc hia howel lii , Trifolium sp., Viola cuneat a , V. ocellata . CHARACTERIS TIC BUTTERFLIES Papilio zelicaon , P. indra spp. (N.W . Cali fornia se gregate ), Euchl oe hya nti s hyant is , Ant hoch aris l ance olata l anceol ata , Coeno nympha calif ornia er yngii , Oeneis nevade nsis id una , Euphy dryas edit ha near colonia , Thessalia leanira n ear orego nensis , Clossiana epit hore c hermoc ki , Speyeria zerene glori osa , Mitoura s pinet orum , M. johns oni , E uphil otes battoi des near orego nensi s , Icaricia s aepiol us ae haja , Hesperia comm a mattoo ni , Thorybes diversus , Pyr gus com munis . DISCUSSION In the O uter C oast Ra nge of t he e xtreme northwe ster n corn er of the state in Del Norte and Humboldt counties, and in adjacent extreme sout hweste rn Ore gon, t here i s a uni que pl ant commu nity on serpe ntine s ubstrat es cha ract erized a s foll ows: dwarfe d indi vid uals of Silver Pine ( Pinus montic ola ) are scatt ered in a n ope n woodl and, wit h inte rsperse d stun ted chapa rral sh rubs a nd areas of spa rse g rass and herb g rowt h on gravel ly to rocky s lopes a nd f lats . There are a lso see ps, sp rings and rivu lets whic h suppor t isl ands of a wet mea dow subco mmunit y withi n the more e xtensi ve dr yish p ortions ; t he boggies t of these w et meadows supp ort popu lations of the unique C alif ornia P itcher - Plant, Darlington ia californica , a nd are si milar t o the b oggy mea dows on ultra basi c subst rates o f the Trinit y Alps re gion desc ribed by Shapi ro et al. (1981). This co mmuni ty is us ually bo rdere d by No rthern Mixed Evergre en Fores t, Douglas F ir Fore st, Lower Montane Conife rous Fo rest, or Mixed Chapar ral. Alt hough it re ceive s high winter rainfal l and freq uent su mmer fog, the shall ow, calcium - poor soil of t he se rpe ntine substrate suppresses vegetation growth in all but the more permanen tly moist portions o f it. The r esult is an admix ture of plants from several commu nitie s in an open s ett ing on t opogra phica lly va ried terrai n, resul ting in nume rous u nique mi croha bit ats and corres pondi ngly a rat her ric h butter fly fauna . This uni que com munity wa s not rec ognize d by Munz (1 959) or Thorne ( 1976). Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 99 29. LOW ER MONTANE CONIF EROUS FO REST DOMIN ANT OR CHARA CTERIS TIC PL ANTS P inus pon derosa , P. jeffreyi , P. coulteri , P. sabi niana , Sequoi adendr on gia gante um , Quer cus k ello ggii , Pse udots uga menziesii , Abi es conc olor , Caloc edrus decurre ms , Pinus lambertiana , Cornus nutta llii ; unders hrubs of va rious s peci es of Arctosta phylos , Ceanot hus , Prunus , Rib es , Chry sotham nus , Eriogon um , and Lupinus , as well as Chryso lepis sempervirens , Chamaebatia fo liolosa , and Rubus parviflorus . On th e transm ontane slopes i n the drier for ests dom inate d by Pinus jeffreyi , common unders hrubs i nclude Artemisia tridentata , Ceanot hus c ordulat us , and Cerc ocarpus ledifolius . BUTTERFLY FOODPLANTS Amorpha c alifor nica , Arabis spp., Arce uthobi um campylopo dum , Artemi sia drac uncul us , Ast ragal us dougl asii , Caloce drus de curre ns , Ceanot hus cor dulat us , C. velutinu s , Elymus glauc us , Eriogonum fa sciulatum , E. kennedyi , E. marifolium , E. nudum , E. wrightii , Ke ckiella lemmonii , Lot us crassifo lius , Lupinus albif rons , L. ande rsoni i , L. formos us , L. latifolius , Pinus jeffreyi , P. ponder osa , Ribes roezlii , Sedum spathulifoliu m , Stre ptant hus be rnardi nus , Symphoric arpos spp., Viola lobata , V. purpurea . CHARACTERIS TIC BUTTERFLIES Papilio ba irdii bairdii , P. eurym edon , Neophas ia menap ia menapia , N. m. teha chapina , Euchloe hyanti s andrews i , Colias occidentalis chry somel as , C. h arfordii , Zerene eurydice , Nymphalis califo rnica , Euphydr yas ch alcedo na mac glasha ni , E. col on col on , E. c. wallacensis , E. c. sperryi , Speyeria zerene zerene , S. z. conc hyliat us , S. coronis semiramis , In cisalia eryphon er yphon , In cisalia mossii h idakupa , I. m . windi , Mitour a nels oni , M. spineto rum , Lycaena a rota aro ta , Glauco psyche piasus pi asus , Euph ilotes battoid es intermedia , E. enopte s enoptes , Icaric ia icar ioide s icari oides , I. i. evius , I. lupini monticola , I. neurona , Ochl odes s ylvan oides sylvanoides , Thorybe s pyl ades . DISCUSSION This pla nt co mmuni ty com bines the subc ommu nitie s of Co ulter Pine Fo rest, Y ellow Pi ne Fores t, and Mi xed Conife r For est. These a re tal l Monta ne Coni ferous Fores ts whic h are do minate d by vario us spe cies of yello w pine: Fore sts d ominate d by Pi nus coulteri are found from the bay area to northern Baja Califo rnia, from the Co ast Ranges southw ard through the Transvers e Ranges a nd P eninsul ar Ra nges, at elevat ions from 1, 000 to 7,000 fe et. Th e fores ts domi nated by Pinus ponder osa and/or P. jeffreyi are found from 1,000 to 9, 500' elevatio n, primar ily in the Sierra Nevada, Klamath Mountain Assemblage, North Coast Ranges , and Tr ansve rse and Pe nins ular Ra nges of southern California. Where these two yellow pine species are sympatric, there is con siderable h ybr idization. On the west slope o f the c entral Sie rra Ne vada, fr om 4,60 0 to 8,4 00' elevation, there tends to be a richer mixture of conifer speci es in addition to the yellow pines, including the Gian t Redwood, Sequoi adendr on gig anteum . Shield s (1966) reported on the butte rfly fauna of a yellow pine forest comm unity i n Tu olumne Count y in t he ce ntral Sierra Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 100 Nevada . His l ist of spe cies showed c onside rable diversi ty, compar able to that found by Emm el and Emmel (1963) in an Upper Monta ne Co nifero us F orest c ommu nity at Donne r Pa ss, Placer County, Californ ia. The high species diversity of this particular commun i ty is due i n part t o its very e xtensi ve ran ge, its diverse f lora, and the numerous microh abitats within it which ac comm odate a wide va riety of speci es. Furt her stu dy of this particular community may justify subdividing it into 4 or 5 communities, each wit h its particu lar floral and fauna l compo sition. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 101 30. UPPER M ONTANE CONIF EROUS F OREST DOMIN ANT OR CHARA CTERIS TIC PL ANTS In the Sierra Nevada, largely dominated by Ab ies magnifica , but often ac compa nied by A. concolor , Pinus c ontor ta mur rayan a , P. monti cola , and Tsuga m ertens iana from the adjacent communities, also with Acer gl abrum , Arctostaphylos nevade nsis , Lon icera conjug ialis , Qu ercus vaccinifolia , species of Ribe s and Symphori car pos , Chrysopsis breweri , and Monar della odorati ssim a . In the Klamath Mountains assemblage, A. magn ifica is fre quent ly acc ompani ed by Abie s amabilis , A. las iocarpa , A. procera , Pinus bal fouri ana , Picea engel mannii , P. breweriana , and Tsuga me rtensi ana . In southe rn Cal iforni a, the commu nity is domina ted by Abi es concol or and Pinus l amberti ana , fr eque ntly incl udes Caloce drus de curre ns , and has a vari ed fl ora of unde rstor y plants , parti cular ly shrub by speci es of Ribes . On drier rock y slopes and ri dges in the Sier ra Ne vada a nd T ransve rse Ra nges, there is an open w oodlan d with Ju niper us occid ental is australis , Cercocarpus ledifolius , Ch rysot hamnus n auseo sus , Holodi scus micro phyllus , Arct ostaphy los pat ula pine toru m , and Pinus jeffreyi . BUTTERFLY FOODPLANTS Arabis spp ., Ceanot hus cor dulat us , C hrysopsis breweri , Cirsi um ander sonii , Collinsia spp., Dicentr a nevade nsis , D. uniflora , Eriogonum marifolium , E. nudum , E. umbellatum , Holodi scus m icrophy llus , Juniper us occi dental is austr alis , Pedicul aris semib arbat a , Penstemon newberryi , Quercus vaccinifo lia , Solidago canadens is , Viol a glabel la , V. purpurea , V. sheltonii . CHARACTERIS TIC BUTTERFLIES Papil io eury medon , Parnassi us clodi us bal dur , P. c. ssp. (Tulare Co.), Euchl oe aus oni des tr ansmont ana , A nthoc haris sara stella , Poly gonia zephy rus , Euphydr yas chalced ona sierra , E. c. ire lan di , E. c. trinitina , E. editha s sp. ( G enera ls Highway ), Chari dryas pal la altas ierra , Charidr yas hoff mann i hoffm anni , C. h. se gregat a , Phyciodes orseis h erlani , Boloria epithore sier ra , Speyeria hy daspe hydaspe , S. h. viridicornis , S. a. ire ne , S. a. dodge i , Mitoura bar ryi s sp. (Sierra Nevada, San Bernardi no Mts.), Eu phil otes batt oides int ermedi a , E. enoptes enoptes , Erynni s proper tius , E. pa cuvius lilius , E. p. callidus . DISCUSSION This montane fores t community is located above the Yellow Pine F orest bu t below t he Su balpine Forest commu nity in the higher mount ain ra nges o f the s tate. T horn (197 6) sub divides this in to 4 commun ities, namely W hite Fir - Sugar Pin e Forest, Red Fir For est, Enriched Con ifer Forest, and Moun tain Junipe r Woodland. Our list of butter fly species and bu tterfly foodplan ts is taken almost entirely from the Red Fir Forest and Enr iched Conifer For est subsectio ns of this plant community . This co mmu nity sho ws butt erfl y specie s dive rsity e quiva lent t o that fo und i n the Lowe r Mont ane C onifer ous Fore st, aga in a result of it s occupa tion of a l arge ge ograp hic are a, the high divers ity o f plant speci es, an d the numer ous mi croha bitats in this comm unity. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 102 31. SUBAL PINE CONIFERO US FOREST DO MIN ANT OR CHARA CTERIS TIC PL ANTS P inus cont orta mu rraya na , P. monticola , P . flexilis , Ts uga mertensiana , Pin us alb icaulis , P. balfour iana; asso ci ated shrubs incl ude Quercus vaccin ifolia , Chrys ole pis sempervirens , Ceanoth us cordul atus , Arctost aphyl os spp. , Ribes cereum , and R. m ontige num . BU TTERFLY FOODPLANTS A stragalus whitney i , Care x spectabilis , Castilleja min iata , Dicentra uniflo ra , Draba spp., Eriogonum incanum , E. nudu m , E. um bella tum , Festuca spp., Lupinus arb ustus , Pteryxia terebi nthi na , Ribes cereum , R. monti genum , Strept anthus tortuos us , Viola pur pure a . CHARACTERIS TIC BUTTERFLIES Papilio zelica on , P . indra indr a , Par nassius c lodius baldur , P. c. near menetries i , Pontia o ccidentalis , Euc hloe hy antis ssp. (Sierra Nevada ), Oeneis chryxus ivallda , Polyg onia ze phyrus , Euphy dryas e dit ha aurilac us , Speyeria egleis e gleis , Satyri um fuliginosum fuliginosum , S. f. m aculat um , Lycaena heteronea submac ulata , Everes amyntula (high ele vation ec otype ), Icaric ia shast a shast a , I. lupini l upini , Hesperia c omm a y osemite , H. nevada occidentalis . D ISCUSSIO N The Su balpine Conife rous Forest i s anot her m ontane forest community which lies above the fir forests but below timberline on the hig her mo untain s in t he sta te. T horn (1976) subdi vides this into 3 communities , namely; Lodgepole Pine Forest, White Bark Pi ne - Mo untain Hemloc k Fores t, an d Limbe r Pine Fore st. Again, for the sa ke of simpl ific ation, we pre fer to us e the all - encompa ssin g com munity of S ubalpi ne C onifer ous Fo rest. These f orests are domi nated by Lodgep ole Pine, Moun tain Hemloc k, or t he vari ous spe ci es of White Pines of timb erline, and ra nge fr om 5, 500 t o 9,50 0' in t he n orther n part of the sta te, and fr om 8,0 00' t o nearly 11,5 00' ele vatio n in t he south ern pa rt of the state. In many areas, this community presents more of an aspect of open woodla nd tha n a de nse f orest, due to the more severe soil and weath er conditions at these elevations, which both stunt the trees and also decrease the po tentially favorable sites for them to grow. As a r esult of this op en woodland feature , there are ma ny shru bs and p erennial herbs scattered on the ope n slo pes an d a varie ty of m icro habit ats are affor ded by the va rious geol ogic s ubstrat es a nd va rying slope e xpos ures. As a c onseq uence t he h umidi ty is rel ative ly hi gh alt houg h it does n ot equal that of t he Uppe r Monta ne o r Lower Montane Conife rous F orest below t his commu nity. S hapir o (197 7) desc ribes the butt erfly fau na of a c ombi ned Subal pine C onifer ous F orest and Arc tic Alpine Cushi on Pl ant Community at Castle Peak, Nevada County , California. The species composition of this particular locality is somewha t less diverse than that of Subalpine Forest and Arc tic Alpine Cushion Plant habita ts to t he sout h in t he hi gher Si erra Ne vada, or to the north west in t he Kla math M ountai ns Asse mblage , proba bly a result of its relative iso lation and non - granitic sub strate. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 109 38. GREAT BASIN SAG EBRUSH SCRUB DOMIN ANT OR CHARA CTERIS TIC PL ANTS Artemisia tridentata , Artemisi a arbusc ula n ova , Atriplex canescens , Ce ratoide s lan ata , Chrysot ham nus nause osus , C. viscidifloru s , Grayi a spi nosa , Pur shia gl andul osa , P . tridenta ta , Tetradymia axillaris , Wyethia mollis , perennia l grasses, and many annuals. BUTTERFLY FOODPLANTS Ar temisia dracu nculus , Astragal us lent iginos us , Castilleja chrom osa , Ceanot hus gre ggii , Chr ysothamnus viscidiflo rus , Colinsia , Eriogonum kearneyi , E. microthecum , E. umbellatum , F estuca spp ., Lupi nus alpe stri s , L. caudat us , L . excubitus , L. inyoe nsis , Machaera nther a cane scens , Pu rshi a glandu losa , P. tridenta ta , Stipa spp., Viola spp. CHARACTERIS TIC BUTTERFLIES Papilio bairdii ho llandi , Colias al exan dra edwardsii , Cercy onis st henel e paul us , C. oetus oetus , Neominoi s ri dingsii pallidus , Euphy dryas ani cia whee leri , E. a. near veazieae , E. editha m onoensis , E. e. fri dayi , Charidry as acas tus acas tus , C. a. vallismortis , Spey eria callip pe neva densis , Apodemia mor mo cythera , Satyrium s aepiu m ssp. (E. slo pe Sierra Ne vada), S. behrii , S. fuli ginos um near semiluna , Lycaena heteronea immacul ata , G laucop syche l ygda mus near colu mbia , G. piasus excubitus , Euphilotes ba ttoides glaucon , E. b. argocy anea , E. rita palles cens , Icar icia icarioides fulla , I. i. eosierra , I. i. argus montan a , I. i. panamint ensis , I. i. near pembin a , I. lupi ni argent ata , Hes peria u ncas gui lianii , H. com ma har palus . DISCUSSION This is an extensiv e semi - dese rt scru b com munity, occu pyin g large areas of the Great Basin, and rather extensive areas within California, primarily east of the S ierran - Cascade Axis. It occupi es dee per, sa ndier s oils of arid s lope s from t he Modoc Plateau a rea southwa rd al ong t he ea st sl ope o f the Sierra Nevada to the Inyo - White Mountains and Death Valley region, southw ard to the higher mo untai ns of the ea stern M ojave de sert and the dr yer port ions of t he highe r mounta ins of sout hern California. The do minant shru b in t his comm unity, Artemisia tridenta ta , is also found i n a numbe r of other semi - desert plant communities , but is he re i n combina tion wi th othe r shr ubs suc h as Chryso thamnus or Purshi a forms a rather d istinctive commu nity. S easona l preci pitat ion in thi s comm unit y ranges from 8 to 15 i nches, s ome o f thi s falli ng as winte r snow, a nd it has a gro wing sea son rangi ng from 3 t o 6 months , depen ding on elevation and latitude. It is fr equently found adj acent to Pinyon - Jun iper Woodland , and fre quently there is con siderable intermixing of the two communities. The butterfly specie s composition of this community is fairly consistent throughout large areas of the Great Basin, and m any specie s or subs pecie s are found primari ly withi n thi s commu nity. The butter fly speci es of this communi ty are largely un ivoltine, with peak fl ight acti vity oc currin g durin g the late spring a nd e arly sum mer months . Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 110 39. MOUNT AIN SAGEBR USH SCRUB DO MIN ANT OR CHARA CTERIS TIC PL ANTS A rtemisi a arbus cula no va , A . tridentata , Eriogonum ovalifolium , Leptoda ctylon pungen s , Are naria kingii glaberescens , Erigeron clokeyi , Hyme noxys cooperi , Lina nthus nuttallii , Koele ria macr anth a , Sitani on h ystrix , an d Stipa pinetor um . BU TTERFLY FOODPLANTS Ca stilleja chromo sa , Chamaebatiara millefo lium , Chrysothamnus viscid iflorus , Eriogonum um bellat um , Lupi nus argent eus te nell us , Stip a pinet orum . C HARACTERISTIC BUTTERF LIES Cer cyonis oetus oe tus , Ne ominois riding sii pallidus , Euphy dryas a nicia w heeler i , Chlosy ne ac astus w aucoba , Callophrys lemberti , Cel astri na argiol us near cinerea , Glauco psyche lygdam us near oro , G. pias us nevad a , Lycaei des melissa frida yi , Ic aricia icari oides albih alos , Hespe ria uncas macsw aini . D ISCUSSIO N This m ontane scrub c omm unity is basic ally a high - altitude versi on of Gre at Basi n Sagebr ush Scr ub, but w ith dif ferin g percentages of dominant plants, as well as a marked decrease in the ove rall height of t he low shr ubs w hich ma ke up t he gr ound cover. Thi s community ranges in ele vation from a bout 8,000 to nearl y 13,0 00 feet , an d is fou nd pri maril y in the Whi te Mountain s, the Inyo Mountains, the Panamint Rang e, the Warner Mount ains, a nd in c ertai n high - elevation localities on the ea st sl ope of the Sierra Neva da. It ha s a sho rt gr owing season and is subject to rat her rigorous weather conditions, includi ng hi gh wi nds, occasi onally dee p wint er sn ows, a nd relatively low seasonal precipitation on the average. I ts butterfly species co mposition is sim ilar to that of the lower elevation, more mesic Great Basin Sagebrush Scrub, but tends to have fewer species o verall. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 111 40. BLACKBR USH SCRUB DOMIN ANT OR CHARA CTERIS TIC PL ANTS Cole ogyne ramosissima ; frequently associated species are Grayi a spin osa , Larrea di varic ata tri dentata , Lycium anders onii , M endora s pinesc ens , and speci es of Ephedr a and Yu cca . BUTTERFLY FOODPLANTS Arabis pulc hra , Castilleja chromosa , Caulant hus coo peri , C. lasiophyllu s , Xylorhiza tortifolia , Yucca species. CHARACTERIS TIC BUTTERFLIES Pontia sisymb rii nigravenosa , Euchl oe hy antis lot ta , Anthoc haris cethur a hadro marmor atus , A. c. moja vensis , A. sara t hoosa , Thessalia leanira alma , Chari dry as acast us neumoe geni , Megathy mus y uccae maudae . DISCUSSION This comm unity consis ts of a low scrub domin ated by Blackbu sh, Coleogin e ramosissima , and i s fou nd on s emi - arid, stony soils at moder ate elevations in both the Death Valley region and eastern Mojave Desert. The butterflies species associated with it also occur in t he adjacent desert plant commu nity. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 112 41. JOSHUA TREE WOO DLAND DO MIN ANT OR CHARA CTERIS TIC PL ANTS Yuc ca brev ifol ia ; frequently associated w ith it are species of Cassia , Eriogonum , Krameria , Lycium , Sal azari a , Tetradymia , Opuntia , and other species of Y uc ca . BU TTERFLY FOODPLANTS Arabis pu lchra , Bouteloua g racilis , Cassia armat a , Ca ulanthus cooperi , C. lasiophyllus , Descurai nea pin nata , Lepidium fremontii , Stanleya pi nnat a , T hamnosma m ontana , Yucca brevfolia . C HARACTERISTIC BUTTERF LIES Papilio polyxenes color , Pontia pr otodic e , P . beckerii , P . sisymbr ia ni grave nosa , Euchloe hya ntis l otta , Anthochar is cethur a mojav ensis , A. sara thoos a , Thessal ia leani ra al ma , Chari dryas a castus neumo egeni , Glauc opsyc he lyg damus desertico la , Euph ilotes rita elvirae , Megathym us yuc cae marti ni , Hespe ria pa haska mar tini . D ISCUSSIO N Th e tre e - like yucca, Y. brevifolia , forms a rather distinctive open wo odlan d in the highe r desert s of th e stat e, from the mounta ins of Joshua Tree Nat ional Mon ument northwa rd to the Owens Valley and Death Valley region. Numerous small shrubs are associated with the Joshua Tree, as well as m any perennia l grasses and nu merous annu als. This community is frequently transitional with oth er desert plan t communities, such as Piny on - Juni per W oodland , Grea t Bas in Sage brush Scrub, Blackbrush Scrub, etc. The butterflies associated with th is community tend to be spring - fliers, but there is a second peak of flight activity in the late summer and early fall months, par ticularly in years of summer rai nfal l. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 113 42. MIXED DESERT SCRUB DO MIN ANT OR CHARA CTERIS TIC PL ANTS Eriogonum fasciculatum polifo lium , La rrea , Ephedr a spp., Ambrosi a du mosa , Atriplex spp., Lycium spp., Prun us fascicula ta , Sphae ralce a ambi gua , Yuc ca spp., Hapl opappus linearifoliu s . BU TTERFLY FOODPLANTS Arabis sp p., Castilleja chromo soma , Caul anthus c ooperi , C. lasiophyllu s , Cymopt eru s panami ntensi s acutif olius , Desc urainea p innata , E riogonum fas ciculatu m , E. pu sillum , Lotus r igid us , Parietaria floridana , Sp haeralcea a mbigua , Thamn osma m ontana , V iguiera deltoid ea , Xylor hiza tortifolia , Yucca s chidi gera . CHARACTERIS TIC BUTTERFLIES Papil io poly xenes color o , P. indr a fordi , Ponti a protodi ce , P. sisymbr ii nigr avenos a , Euchloe hy antis l otta , Anthoc haris cethura near ha dromar morat us , A. c. mojavensis , C hlos yne californ ica , Charidry as acast us neumo egeni , Euphilotes battoi des m arti ni , E. mojave , Megathy mus y uccae maudae , Pyrgus albescens , Heliopetes ericetorum . D ISCUSSIO N This nam e we have coine d for a rat her exte nsive desert sc rub commu nity wh ich is usuall y lum ped wit h Cre osote B ush Sc rub. It is p robabl y simi lar to t he c oncept of Tho rn's (1976) non - basic rock pla nt subc ommu nity of h is Dese rt Rock Pl ant comm unit y. But we conside r it to c over mo re ext ensive habit ats. As concei ved h ere, t his com munit y consi sts o f a low scrubl and commu nity, primari ly occ upying sl opes of the des ert mounta ins, a bove t he Cre osot e Bush Sc rub Com munit y of t he flatl ands and v alle y bottoms , but belo w the othe r desert pl ant commu nties such a s Bl ackbus h Scrub , Jos hua Tree Woodl and, and Pinyon - Juni per Woo dland. Creosot e Bush ( Larre a ) is usual ly prese nt i n this comm unity, but te nds to occur only a s scatt ered pl ants, in cont rast t o the " dense r" sta nds of t his spe cies on the flatl ands below . Thi s comm unity occupi es l arge a reas o f the Mojave Desert, as well as some of the l ower elevatio ns of the Death Valley region, the east slopes of the Sierra Nevada in Inyo C ounty, and t he hig her e levat ions o f the Colo rad o Desert. T he butterfly species composition of this community is similar to that of the other desert plant co mmunities. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 114 43. CALCICOLOU S DESERT SCRUB DOMIN ANT OR CHARA CTERIS TIC PL ANTS Agave utahensis nevadens is , Ar temisia bigelovii , Buddleia utahens is , Cercoc arpus i ntri catus , Fendlerella utah ensis, Forsellesia nevadensis , Pe trophyt um cae spit osum , an d Symphor icar pos l ongifl orus , and such her bs as Arab is perenn ans , Brickellia arguta , C ymopt eris aborigi num , Eriog onum i ntrafr actum , Hedeo ma nan a califor nica , Hofmeistera p luriseta , Mauran dya pet rophil a , Parietaria floridana , Penst emon c alcare us , Phacelia rotundifolia , Salvia funere a , and Scophuloph ila rixfordii . BUTTERFLY FOODPLANTS Aga ve deserti , A. ut ahens is , Arabis pe renna ns , Castilleja chrom osa , Caulant hus gl aucus , Eriogonum h eermannii , Maurandya antirr hiniflora , Pet rophyt um ca espitos um , Pursh ia stansbu riana , Thamnos ma mont ana , Yucca schidigera . CHARACTERIS TIC BUTTERFLIES Papil io poly xenes color o , Pontia sis ymbrii nigrav enosa , Euchl oe hyant is lott a , Thessal ia leani ra alma , In cisalia fotis mojavensis , Call ophrys coms toki , Celastr ina argiolu s near cinerea , Eup hilot es batt oide s eurom ojave nsis , Megathy mus yucca e maudae , Agathy mus al liaea ssp., A. baueri ssp. DISCUSSION This desert scrub community is similar in appearance to the Mixed Deser t Scru b comm unity , but has a diffe rent p lant species composition as a result of its location on Calcar ious rock subs trate s (lime stones , marbl es, and dol omites ). Many of the plants of t his com muni ty are c onfi ned to calca rious substrates, but deser t species from oth er soil substr ates are also found mixed with this flora. It is found in scattered localities prima rily i n the e aste rn Moja ve and Deat h Valle y regio n mounta in range s, from a ppr oximatel y 3,0 00 to 8, 000 f eet elevation. At least one butterfly species ( Agathymus alliae ) is confined to this community due to the restriction of its foodpl ant t o cal carious subs trate s, and severa l ot her de sert butterfly species are found primarily in this plant community as a res ult o f the ir foo dplant s growi ng i n abun dance on t his particular sub strate. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 115 44. CREOSO TE BUSH SCRUB DO MIN ANT OR CHARA CTERIS TIC PL ANTS Lar rea div aricat a trident ata ; often ac compa nied by Ambrosia dumosa , Atriplex spp., Ce ratoides lanata , D al ea spp ., Encelia farinosa , Eriogonum inflatum , Fouq uieria splen dens , Gra yia spinos a , Hymenocl ea s alsola , Lepidium fremontii , Lycium spp., Menod ora spi nesce ns , Sphaeralc ea ambigua , along w ith scattered Yuccas and various cacti of the genra Ec hinocer eus , Feroc actus , and Opuntia . Also annua ls and perennia ls in the genera Abron ia , As tragalus , Camissonia , Chaenactis , Choriz anthe , Coreopsi s , Cry ptant ha , Eriogonum , Erioph yllum , Gilia , Linant hus , Malacothr ix , Men tzelia , Nama , an d Oenothe ra . BUTTERFLY FOODPLANTS Am sinckia spp. , Asclepias er osa , Caul anthus cooper i , C. lasiophyllu s , Cleo me isomeris , Cryptantha spp. , Descura inea pinnat a , Eriogonum fasciculatum polifolium , E. in flatum , E. pusillum , E. ren ifo rme , Geraea canescens , Hilaria rigida , Lepidi um fre monti , a nnual Lupi nus spp., Pala foxia linearis , Sphaer alcea a mbigu a , Streptanthella longirostr is , Vi guier a deltoidea , Xylorh iza tortifolia . C HARACTERISTIC BUTTERF LIES Pont ia prot odice , P. beckerii , Euch loe hyantis lotta , Anthoc haris ce thura ha droma rmorat us , A. c. mojave nsis , A. c. pima , Nathalis io le , D anaus ple xippus , D. gilippus , Van essa cardui , V. annabella , C hlosy ne cal ifor nica , Charidry as acast us neumoe geni , Apodemi a mormo de serti , Euphi lotes b att oides martinia , E. mo jave , Copaeo des aur antiac a , Pyrgus albescens . DISCUSSION On the valley floors, f latlands, and bajadas of th e California desert s, fr om nea r sea le vel u p to ap proxi matel y 4500' elevation, th ere is an extensive scrub community domin ated by the s hrub, L arrea di varic ata tride ntata . Although Larrea is the domina nt pla nt of t his c ommuni ty, t here a re num erous small shrubs growing with it, as well as many smaller perennials and numer ous ann uals w hich, in years of favora ble wint er rain fall, can bec ome a m ass of c olor i n an exte nsive s pring bl oom. Follo wi ng su mmer rains, there may also be a similar but less specta cula r bloom of su mmer annua ls, w hich a re o f diffe rent species th an those triggered by winter rains. Mos t rainfall in this c ommuni ty fa lls dur ing t he winte r mon ths, a nd even then tends to be vari able a n d unrel iable fro m year to year. In s ome dry yea rs, virt uall y no prec ipit ation w hatsoe ver fal ls, an d that partic ular sea son ma y be devoid o f butter fly speci es. Wint ers are c ool, fr equent ly win dy, a nd sum mers t end t o be ver y hot and punctua ted b y only rar e t hunders torms. N ot surprisingly , virtually all bu tterfly flight ac tivity occur s durin g the spri ng mo nths, wi th a pea k in la te Marc h or ea rly April. For a few species of this community, a late summer or fall br ood ma y be t rigge red by summe r rai ns. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 116 45. SEMI - SUCCUL ENT SCRUB DO MIN ANT OR CHARA CTERIS TIC PL ANTS Agave deserti , Nolina s pp., Yucca sc hidi gera , Feroc actus acanthod es , Fougui eria s plende ns , Larrea , Enceli a farin osa . BU TTERFLY FOODPLANTS Aga ve deserti , Yucc a schidi ger a . C HARACTERISTIC BUTTERF LIES Megathymus yuccae ssp. ( wester n edge Col orado Des ert), Agathy mus st ephe nsi . D ISCUSSIO N This desert scrub c ommu nity oc curs prima rily on the r ocky upland slope s of the dese rt mo untain s in the Colora do D esert, and is domina ted by s emi - succul ent spe cies in the ge nera Agave , Nolina , and Yucca . It is more likely to receive summer rain, and in fact appr oximately h alf of the rather limited ann ual preci pitat ion in t his c ommuni ty falls d urin g the s ummer months . The nu mber o f butte rfly s pecie s found i n this community is rath er limited, with two of the most characteristic skipper species us ing t he A gave and Yucca species as foodpl ants. One of the se sp ecies is a s pring - flier, while the other is a fall - flier, and neither seems to be influenced greatly by differences in annual precipitation , in contrast to virtually all other des ert butterfly species. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 117 46. DESERT MICROPHYL L WOO DLAND DO MIN ANT OR CHARA CTERIS TIC PL ANTS A cacia gre ggii , Cercidium floridu m , Dalea spi nosa , Olneya tesota , P rosopi s glan dulosa t orre yana , P . pubesc ens , Chilopsis linearis , H yptis emoryi , and Larrea . Smal ler sh rubs incl ude Calliandra erioph yllum, Cassia armata , Hoffma nnseggi a microphylla , Bacc haris s arothr oides , Condalia lyvio ides , Hymenoclea monogyra , an d Lycium a nderso nii . BU TTERFLY FOODPLANTS Acacia greggi i , Asclep ias subulata , Bebbia j uncea , Calliandra eriophyllum , Cassi a armata , Hibi scus den udatus , Hoffm annseg gia mic rophy lla , Justicia californica , Palafoxia linearis , Phorade ndro n cali fornicu m , Proso pis gland ulosa . CHARACTERIS TIC BUTTERFLIES Eurema ni cippe , Nathalis iole , Da naus gili ppus , Dym as ia dymas i mperi alis , Calephelis wrighti , Min istrymon led a , Atlides halesus , Strymon co lumella , Lept otes mar ina , Hemiar gus isol a alce , H. cera unus gyas , S ystasea zampa , Erynnis funeralis . D ISCUSSIO N This c ommu nity is c hara cteri zed by conce ntrati ons o f Fabace ous des ert shr ubs loc ated pri mari ly alon g washe s and other desert wat ercours es or drai nagewa ys, a nd is f ound primarily in the inter - mountain valley s of both Calif ornia deserts , prima rily below 5,00 0' elevation. Virtually all th e butterfly species associated with th is community are also fou nd in the sub tropical thorn fo rests of Mex ico, and repre sent a northe rn ext ensi on of t he ra nges of those speci es. Some of th ese specie s may n ot be re sident in ma ny of the desert localities in which t hey ha ve been r ecor ded; c onsidera ble fl uctuat ion in abunda nce occ urs f rom ye ar to ye ar, an d undou btedl y some popul ations are rei ntro duced b y migra nts from m ore pe rmanent popul ations to the sout h. There is n o high c onc entration of flight activity in one particular season of the year as there is with t he o ften adjacent Cre osote Bush Scru b plant comm unit y butterfly sp ecies; due to the variations in blooming times of some of the dif ferent butt erfly fo odplant s of t his com mu nity, as well as variatio ns in winter and summer precipitatio n, as well as cons idera ble fluctua tions in nu mbers of im migra nts fr om population s outside the area, no distinctive pattern of flig ht activity seems apparent from year to year. In drier years, mu ch of the flight acti vity se ems t o conce ntrate in t he lat e spring and early summ er months, whe reas in years w ith heavy summer rains, there ma y be inc reasi ng abu ndance o f speci es throu ghout the su mmer m onths with a peak a bundance reac hed in t he fal l months . Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 118 47. DESERT RIPARIA N WOODLAND DOMIN ANT OR CHARA CTERIS TIC PL ANTS P latanus race mosa (of western deser t margin s), P opulus fremontii , Salix gooddingii , S. laevigata , F raxinus velut ina , Bacc haris ser gil oides , Washi ngtoni a fil ifer a , Pluch ea sericea , Phragmites a ustralis , A rundo donax , and species of Typha an d Bacc haris . BUTTERFLY FOODPLANTS Atrip lex canescens , Bacchari s glut inosa , Phoradendron californ icum , P. flavescens macrophyllum , Populus f remont ii , Prosopi s gla ndulos a , P. pubesc ens , Salix spp ., Sesba nia exaltata . CHARACTERIS TIC BUTTERFLIES Limenitis a rchippus obsoleta , Apodem ia palmer ii , Ca lephelis nemesis dammersi , Atlides halesus , Plebul ina e migdi onis , Erynnis funeralis . DISCUSSION This desert wood land community is found primarily along the Colora do an d Moj ave R ivers , an d to a less er exte nt ne ar t he Salton Sea and alo ng irrigation canals in the Imperial Valley. Numer ous but terfl y speci es, many as s trays , ha ve been t ake n in this type of habitat, bu t are not "nativ e" to it, having been attracted to it an d concentrated around it pr imarily by the presence of water and therefore nectar sources. However, there are a small number of species which seem regularly associated with it and ar e dependent on foodplants requir ing this habitat. Much of the ripari an w oodla nd alo ng t he Colo rado R iver has been gre atly disturbe d by vari ous da m proje cts and other extensive changes along that particular watercourse, and much of the ve getat ion which is presently repr esented there is the result of co nsidera ble m an - m ade dist urbanc es. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 125 On the eastern side of the state, bo rdering Nevada, lies the m ajestic Sierra Nevada Range, extending north - south for some 430 miles. The east ern escarpment of this range is sheer, dropping some 10,000 feet within ten miles near Owens Lake. On the wester n side, however, the range slopes more gently to the Central Vall ey in gradually declining foothills. Arising near volcanic Lassen Peak in the north, the Sierra Nevada extends south to the fringes of Los Angeles. It is approximately 50 - 80 mil es in width and contains about 27,000 square mil es of area in its 430 mile length. Mt. Whitne y, the highest point in the continental United Stat es, and ten other peaks exceed 14,000 feet in altitude. Very few p asses cross from east to west over th e Sierras and sever al of these occur at more than 9,00 0 feet elevation. The largest lake in the Sierr a Nevada is Lake Tahoe, extending across the California - Nevada border at 6,229 feet above sea level. It is almost 200 square miles in area, and is remarkably deep, with the maxim um depth exceeding 1,600 feet. Throughout the Sierra lie hundreds of smaller lakes, many above timberline in regions of tumbled granite and smooth walled canyons. There are three great national parks in these mountains: Kings Canyon, Sequoia, and Yosemit e, the las t being part icularly famous for it s glacie r - carved valleys of the Merced and Tuolumne Rivers, with their many waterfalls and granite domes. Central Valley Between the coast al ranges and the Sierra Neva da, the great C entral Valley runs fo r 450 miles in a north- south arc. Once a paradise of flowers and natural grassland, with scattered oaks, today the Valley constitutes the sta te's agricultural heart land. I ts sing le exit is the delta thro ugh which the Sacramento and San Joaquin Rivers drain into San Francisco Bay. The northeastern edge is bordered by the Cascade Range and the northwestern edge by the Klamath Mountains. In these northern areas, few people live and the terrain is quite rugged and heavily timbered. The coastal sides of the Cascade an d Klamath Mountains tend to be w et, while the northeastern side s are drie r and more barren o f vegetati on. At th e southern end, th e Centra l Valley is closed off b y the several T ransverse Ranges, esp ecially the Teh achapi Mountains. The Tehachapis are regarded as a dividing line w all between south ern and central Califo rnia. Deserts The deserts of east ern and southeastern Cal ifornia are quite var ied in vegetation and topography. The Colorado Desert of southeastern California has summer temperatures of up to 130 o F (54 o C ), with annual ra infall a veraging 3- 4 inches. Much of this desert area lies below sea level. The i nland Salton S ea (300 square miles in area) is entirely be low sea level and was creat ed in 1905 - 1907 when the Colorado River broke out of its channels and poured into an interior desert basin with no outlet. Northward, in t he higher Mojave Desert, which can range 3,000 to 4,000 fee t on the desert fl oor and up to 7,000 feet or so above sea l evel in the mountain ranges there, temperatures are so mewhat less e xtr eme. This area of more than 25,000 square miles reaches from the Tehachapi and the Sierra Nevada to t he Colorado River, and is still relatively untouched by developm ent. Min ing comp anies, mil itary re servation s and avia tion tes t facilitie s have dis turb ed some areas. The eastern G reat Basin Desert area lying in the eastern ra inshadow of the Sierra Nevada Range rises from 2,000 to 7,400 feet above sea l evel and has temperatures ranging from 75 o F (24 o C) in the summe r to -25 o F ( -32 o C) in the w inter. All of the se geog raphic reg ions o f the state collectiv ely repr esent a varied n atural landscape, one in which butterflies of all groups h ave found hospitable niches to inhabit and evolve Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 126 new and marvelous forms that we find today. The unique geographic diversity of California has thus contributed directly towards generating the exceptional butterfly species diversity found in the state. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 127 ORIGI NS OF THE CALIFO RNIA BUTTERF LY FAUNA ORIGI NS OF HOS TPLANT FLO RAL ELEMENTS The origin s of the Ca lifornia bu tterfly fauna are r elated in sub stantial p art to the geo flora sources generating the d iverse plant assemb lages found in the state. First, as noted in the precedin g chapter, many rel ic plant g roups from the Tertiary Period (63 t o 2 million years ago) of the Cenozoic Era have survived in the California Floristic Province because of the mild climate o f the region and its r elative i nsulation f rom the ex tremes of the continen tal clima te for the past mi llion years o r so. A dditio nally, there has been a substantial interplay betw een the mesic (mo isture - lovin g) flora l elements derived fro m the Arct o-Tertiary G eoflora w ith the xer ic (dry - loving ) floral e lements assoc iated with the Madro -Terti ary Geoflo ra. With th e great topographic relief and varied soil conditions in this region, many new combinations of plants could evolve in new community relationships. Third, some of the more important genera that are butterfly foodplants, such as Lupinus and Penstemon , have shown substantial bu rsts of speciatio n and clusters of speci es have been pre served in limited areas of the region. S ome of the plants (about 15% of the specie s in the Ca lifornia F loristic P rovince ) appear to h ave radiated into our state fro m the deser ts of other, more southe rn parts of N orth Americ a (e.g. , species of the family Malvaceae). Many fam ilies of flowering pl ants seem to have co me in an ultimate sen se from South America to North America (Raven and Axelrod, 1974: 627-628; Raven, 1988). These include many of the leg umes and such families as the Lauraceae (host s of some of our swallowta ils) and A steraceae (h osts of a number o f our nympha lids). Add itionally, there are some similarities ex isting between th e floras of the Medi terranean region and Californ ia, which in part have had their origins via differentiation from a common northern flora, including such important b utterfly ho st genera as Juniperus , Populus , Prunus , Rhamnus , and even Pinus and Quercus . Nonetheless, the unique character of the flora o f California has been st ressed by many workers. C onsider that, within the Californ ia Floristi c Province alo ne, some 58 g enera (7.5% o f the total genera) and 2133 of the 4427 species (48%) are en demic. However, taking a broader view and lo oking at the u ltimate sou rces of the f lora, Raven (1988) fee ls that "it is c lear tha t the flora of C alifornia can be rega rded as c omprising two element s: a northe rn, or A rcto - Tertiary, and a southern, or Madro- Tertiary, element." Because the Medi terranean climate had de veloped after the mid- Pl iocene (see Chapter 2 ) in the broad eco tone area occu rring between these fundamentally differen t vegetation ty pes, a furthe r stimulus fo r the prolifer ation of spe cies and even so me ge nera was present. During relativ ely recen t times (the la st 10,000 y ears), additio nal plants f rom the drier parts of the Madro - Terti ary region hav e entered the Californ ia Florist ic Provin ce and especia lly colonize d the drier v alleys o f cismontan e to sou the rn California, the San Joaquin Valley, and the inner South Coast Ranges, evolving a number of endemic species in that region. But the influence s on the Californi a flora are not rest ricted so lely to tho se contrib uting to the origin s of plants within th e Californ ia Flori stic Provinc e. After all, while that provin ce comprises about 75% of the state's area (with some 744 of the 875 genera [85%] and 4,119 of the 5,057 species [81%] found in the s tate), we also have a large number of genera (approximately 131) and 938 species that occur onl y east of the Sierra Nevada mountains and on the deserts, but still within the boundaries of California. Stebbins and Major (1965) have pointed out that among 10 biotic subdiv isions st atewide, there are four distinct tr ansmontan e reg ions in th e eastern and southeas tern portio ns of C alifornia with affin ities outsid e the state. The se are shown in F ig ure 5 - Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 128 1. These vegetational a reas each h ave had an impact on the in fluence of e xtra - Califor nia butte rfly faunal regions on the state's butterfly fauna, again because of the i ntroduction of larval foodplant elements. Great Basin Region The Great Basin Region occupies the nort heastern corner of California, including all of Modoc County and portions of Lassen, Shas ta, and Siskiyou counties. It also i ncludes the northernmost part of Inyo County above the Owens Valley, and Mono County east of the Sie rra Nevada. The flora of this vast region, comprising approximately 22,600 km 2 , is essentia lly that of the Great Basin, although a number of taxa of the California flora extend beyond the Sierr a- Cascade axis to the Warn er Mountains and other portions of the Modoc Plateau. W hile there are only two endemic plants re stricted to the n orthern p art of th is Great B asin regio n withi n Californi a ( Lomatium ravenii and Mimulus pygmaeus ), there ar e at least a dozen or so other endemic species in the southern part. Additionally, both the northern and t he southern parts of the Great Basin region in California have over 120 and 147 specie s, respectively, that do not occur in the California Floristic Province, including 67 and 26 species each that are found no where else in the state. The White Mountains, situated at the southwestern corner of the Great Basin region, contains some 763 native species a nd varieties (Lloyd and Mitchell, 19 73). These latter authors conclude that the White Mou ntains are a Great Basin Desert r ange with a f lo ra enriched during the Pleistocene by many boreal elements v ia the Sierra Nevada Rang e. These are especially fou nd in alpine and subalpine a reas, while other families s uch as Fagaceae and Ericaceae a re absent in the White Mountains. From the south have come various Sonoran and Mojavean taxa. Inyo Region The Inyo Region is, as Raven (1988) points out, the richest and m ost interesting floral region in transmontane California. Here ar e found some 454 plant species that onl y occur within this region in California, including at least 43 endemic species. Eve n some genera are en demic in the Inyo Region, including those in the Death Valley a rea. Some of the butterfly foodplants and nectar sources occurring in California only in this region include Buddleia , Cowania , Enceliopsis , and Monarda . Endemi c plants developing on limestone in the deep canyons f lanking Death Valley are a part icularly imp ortant fe ature o f the Inyo R egion. Mojave Desert Region In the Mojave Desert occur about 757 plant species, of which 22 are endemic. Over 50 of the species in the Mojave Desert are not found in any other part of the st ate, but range into Nevada or northwestern Arizona. Approximately half o f the plant species in the Mojave Desert extend westward to the mountains of southern Calif ornia, the San Joaquin Valley, or the inner South Coast Ranges, and thus occur in the California Floristic Province as well. Sonoran Desert Region The Colorado Desert represents the northwestern arm of t he Sonoran Desert, and has about 306 species not found in the California Floristic Province. At l east 100 species are included that do not occur elsewhere in the stat e, and there are at least eight end emic plant species. Th e Colorado Desert recei ves summer rain sto rms, whereas much of the Mojave Desert lacks summer rain, and this facto r may restr ict a numbe r of desert p lants in th e Californi a portion o f their ra nge to the Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 129 Colorado Desert. Accordingly, the butterfly faunas of the Color ado Desert and the Mojave Desert are substa ntially differen t, althou gh some b utterfl y species range north ward to the Mojav e Desert or even to the Great Basin deserts of Inyo County as well. ORIGI NS OF CALIF ORNIA BUTTE RFLY FAUN AL ELEM ENTS Palearctic Influen ce The butterfl y fauna of California co ntains a sizea ble number of spec ies that are very closely related to European butterflies. Many genera and even so me species are held in common. For example, the modern genera shared by Europe, northern Asia, and North America include Papilio , Parnassius , Colias , Pieris , Coenonympha , Oeneis , Vanessa, Nymphalis, Polygonia, Lycaena, Ochlodes, Erynnis, and Hesperia . We could easily expand this list to include genera which are likely identical, su ch as Argynnis / Speyeria an d many of the lycaenid groups. The extraordinary similarity among man y of the p arnassians, friti llaries, rin glets, co ppers, hairstreak s and blues is related to the physic al conn ection o f the Nea rtic (North America ) and Palear tic (Eu rope and northern Asia) regions across the North Pacific by the Bering l and bridge and the Aleutian arc islands. All these a reas were expose d during Pliocen e and Pleistocene periods of low er sea level and heigh tened gla ciation , and butt erflies w ere able to cross from Sib eria to A laska and proceed south down the Cascades and Sierr a Nevada Range. The southern lim it was reached about the level of the Kern R iver and Walker Pass. Presumably, man y of these species occur red at lower elevations at such times, as glaciers covered much of the high country in the Sierra and the cool er temperatures would have forced hospita ble vegetation zones for these northern species lower on the mountains. Great Basin Influence Another larg e element of Californ ia's butte rfly fauna penetrate s from the Great B asin are as. These includ e many checkerspots o f the genera Euphydryas and Chlosyne ( sensu lato ), Cercyonis woodnymphs, the admiral Limenit is weidemeyerii , skippers in the genus Hesperia , and blues such as Icaric ia icarioides and Lycaeides melissa . These deser t - adapted species h ave frequently undergone phenotypic divergence in both the Great Basin mountain ranges and the elevated desert ranges in Ca lifornia. Southern Desert In fluence Further to t he south, a substantial po rtion of the b utterflies current ly in Califo rnia came in from the Sonoran Desert areas of Arizona and northern Mexico, having foodplant members of the Madro - Terti ary flora. These butterfl ies, characteristic of desert , grassland, open w oodland, and even thorn forests, pen etrated th e desert reg ions of south ern and east ern Califo rnia, from th e Colorado River west to the cismontane reg ion. Notable faunal elements for the but terflies include the family o f giant skip pers ( Megathymidae). Except for several coastal and interior mountain populations in San Diego County, the megathymids are largely restricted to the Mojave and Colorado d eserts in California . Here , too, are found most of the Californ ia metal marks of th e family Riodinidae. Only the Mormon Metalmark ( Apodemia mormo ) extends its range as far as the San Francisco Bay area of norther n California. Neotropical Influen ce A fourth influenc e on bu tterflies has been the inv asion of n eotropical species, follow ing the introduction of subtropical and tropical plants into Cal ifornia for horticultural use. Our only Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 130 representa tive of the subfamily Heliconi inae, the G ulf Fritill ary ( Agraulis vanilla e ), has successfully estab lished itself becau se of widespr ead plantings of Pass ionvines (genus Passiflora ) in landscaping, from the southern border of Cali fornia to as far north as Redding i n the Sacramento Valley. The plan ting of citrus tree s from so utheas tern Cali fornia and the Imper ial Vall ey, up int o the San Joaquin V alley at least as far north as Po rter ville in ce ntral Cali fornia, has m ade possib le the expanding introduction of the Giant Swallowtail ( Papi lio cresphontes ). Likewise, the plantin g of Cas sia shrubs in southern California north to San Jose an d Santa Clara County have allowed the range expansion of the Cloudless Sulphur ( Phoebis s ennae ) and the Orange Sulphur ( P. agarithe ) from more tropical cli mes. Loss of Floral Influen ces Concomitan tly, the lo ss of floral e lements in certain areas has resulted in extinctio n of loca l butterfly fauna. San Francisco on the northern California coast has been extensively surveyed b y botanists since th e time of the foun ding of th e local P residi o and Mission in 1776. Here was a unique mingling of the north coastal and south interior elements, and a small but impressive group of endemics found nowhere else. Up to 1940, the natural wild areas i n San Francisco seemed relative ly secure fro m human d estruction, but after Wo rld War II, vast housing pr ojects and industrial developments marched up even the steep hillsides of the San Francisco peninsula and much of the flor al diversity began to disappear (Howell, Raven, and Rubtzoff, 1958). Far ea rlier t han this, the unique sand dunes areas and their vegetation that supported such unusual endemic butterflies as the Boisduval's Satyr ( Cercyonis sthenele sthenele ) and the Xerces Blue ( Glaucopsyche xerces ) were consumed by the expanding urban areas, and thes e butterflies became extinct by 1880 and 1941, respectively. Today, other characteristic San Franc isco butterflies are going the same way, including the San Bruno Hairstreak ( Incisalia mossii bayensis ) and the Bay Area Checkerspot ( Euphydryas editha edit ha ). The unique mild coastal climate of these sand dunes and the shrouded hillsides of the California coastl ine have given rise to many en demic butterfly sp ecies and subspeci es. Some are now gone. Others remain, such as the r inglet, Coenonympha ampelos yontocket , restricted to a remarkably small a rea amo ng the co astal sand dunes n orth of C rescent C ity in Del Norte Cou nty. Considera tion of suc h restricted patter ns of distrib ution and en demism is s ufficiently important to be treated in a section of its own. CENTERS O F PLANT ENDEMI SM I N CALIFORNIA As in the ov erall dis tribution s of butt erfly and pla nt diversity in Califo rnia, the c enters of endemism for but terflies in the st ate relate in pa rt to cen ters of e ndemism for plant s. Stebbin s and Major (1965) and Raven (1988) provide excellent summaries of plan t endemism i n Californ ia, from which this overview is taken. One group of endemic species are the Paleoendemics . These plan t species are found in relic are as and are the s ole rep resenta tives of the ir genera in C aliforni a or are sepa rated fro m other species of th eir genera by conside rable geographic d istances. The distribu tion of these r elic species is shown in Figure 5 - 2. Two areas o f maximum concentrat ion of relic plan t species are obvious. One is in the Klamath Mountain area of northern Californ ia, adjacent to southwestern Oregon, where Arcto - Te rtiary relic pl ants pr edominate. The o ther center for relic sp ecies is along the Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 131 northern and western margin of the Colorado Desert, from the Litt le San B ernardino Mo untains along the east slop e of the San Jac into and Santa Rosa Mountains, the Borrego Val ley area, and southward into Ba ja Califo rnia, whe re Madro- T ertiary relics abound. Axelrod (1966) points out that these two regio ns r epresent areas in which relic plant sp ecies and genera asso ciated with the Arcto- Tertia ry and Madro - Tertiary geofloras hav e persisted. T hus, as Raven (1988) states, "It h as been the interplay between thes e elements form the Miocene onward that has given the flora of California its richne ss. Between the m, a region of rather low frequency o f relic species, compar ed with the diversi ty of th e flora as a wh ole, exte nds from the inne r South C oast Ra nges to th e Tehachapi Mountains. This is an area of ac tive speciation but not of relic endemism. " The other areas of richest endemism in the state identified by Stebbins and Major (1965) in sampling endemic species found in 70 large genera of the California flora. As the numbers on the map in Figure 5 - 1 sh ow, the areas rich est in endemic s in these ge ne ra are Southern California , the Sierra N evada , and the Central Coast . The poorest areas are the C entral Valley and the d esert. A high degree of end emism seems t o occur in areas that hav e a great var iety of habitats tod ay and that, during t he Pleistocen e, at least, were not subject ed to such cat astrophic changes as glacia tion. More endemic plants are found in the low mountain areas than in the very high mountains (Stebbins and Major, 1965; Raven, 1988). Many of the endemics that have developed in California have done so in association with unique edaphic conditions. Thu s the restri ction of pl ant taxa t o certain soi l types is an important fe ature of th e flora in the state of C alifornia, and this may be as commonplace as sand dunes along t he coast to podsoils in the Pigmy F orest area of Mendocino County or the serpentine and limestone rock regions of the state. CENTERS O F BUTTERFLY E NDEMISM IN THE STAT E Butterfly endemism in general is associa ted with th e cent ers of plan t endem ism in Californ ia. The se are more f inely d efined, ho wever, by the fact th at butte rfly spec ies norma lly tend to be less widely distributed than their hosts. Thus the Veined Blue, Icaricia neurona , is restricte d to severa l localiti es in the western Tra nsverse Ra nges of So uthern Cali fornia, w hile its larval host, Eriogonum wright ii , is a very widely distributed plant. Just to the north of these Ranges occurs another rare ende mic blue , Plubulina emigdionis , known only from a few colonies; yet its larval host plant, Atriplex canescens , enjoys a vast di stribution throughout the west. We identify seven centers of butt erfly endemism in California. (1.) First, the San Francisco Peninsula had several endemic spec ies and subsp ecies restricted to the sand d une areas and interior hills along the coast, including C ercyonis sthenele sthenele (extinct in 1880), Glaucop syche xerces (extinct in 1941), Icaricia icarioides pheres (extinct in San Francisco), and Parnassius clodius strohebeeni (las t recorded in 195 8 from the moist forest areas near the base of the peninsula). (2.) The second center of endemism was the El Segundo Sand Dunes and Palos Verdes Peninsula area o f the Los Angeles basin. H ere once occurr ed an extensive sand dun e habitat area and coastal mountain - beach interface w hich supported such species as Euphilotes b attoide s ally ni and G laucopsyche ly gdamus palosverdensis . At the opposite end of the state in Del Norte and Humboldt counties, and also along the coastline, occur such r arities as the ringlet Coenonympha Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 132 ampelos yontocket , a relic hairstreak population recently named Inc isalia polios maritima , and the beautiful d ark fri tillary S peyeria hipp olyta (3.) A thi rd center o f different iation is in the Tr ansverse Ranges of southern California. The San Bernardino Mountains represent t he site of probable origin for the beautiful California Dogface, Zerene eu rydi ce , splitting o ff in the distan t past fro m the lowl and and much more widel y distributed North American Dogface, Zerene cesonia . The adaptations associated with movement into the cooler mountains and foothills of these inland montane regions may have provided the opportunity for the evolution of thi s ir idescent o verlaid p ierid. A dditionally, Colias harfordii, Euchloe hyantis andrewsi, hairstreaks su ch as Incisalia mossii hidakupa , Mitoura thornei, Mitour a barryi subsp ecies, and Mitoura loki , as well as metalmark subspecies of Apodemia mormo at various elevations, blues such as Glaucopsyche piasus gabrielinus, Euphilotes battoi des vernalis , all attest to t his area's interest as a c enter of spec iation and end emism. (4.) The fourth cen ter of en demism and evolutio n for Cal ifornia b utterfli es has bee n the desert mountain ranges of both the M ojave and Colorado deserts . Virtually every range has its own distinctive subspecies of Euphydryas chalcedona . A re markable number of sub species and well diffe rentiate d populat ions of P apilio indra have developed in these ranges. Many blues and other lycaenids have evolved distinctive subspecies here. (5.) A fifth center is the Death Valley region of C alifornia. Here, distin ctive subspecies of Papilio ind ra have evolved, along with various desert checkerspots and blues, such as Chlosyne acastus vallismortis and Icaricia icarioides pani mentinus . (6.) To the north, in the alpine regions of the White Mountains and Sierra Nevada Range , we find a six th center wh ere addition al endemic species and subspecies have evolved. Perhaps the most notable is the green - colored sulphur, Colias behrii , found nowhere else in the world except in the alpine areas of the highest part of the Si erra. Additionally, the remarkable new species o f alpine blue, Agriades cassiope , is found nowhere else bu t in the central and northern Sierra Nevada. The beautiful copper, Lyc aen a cupreus periglenes , i s found only on the highest summits of the White Mountains and the Sierra Nevada. Finally, w e should no t fail to men tion the no rtheastern corner of Ca liforn ia as a sev enth center of endemism. Influenced by the Great Basin species fauna but also having unique endemics of its own, the northeastern montane fauna includes the unusual copp er s Lycaena nivalis and Lycaena gorgon jacquelinae , and the checkerspot Euphydryas editha warnermontana . PRESENT PA TTERNS OF BUT TERFLY DISTRI BUTION A ND DIVERSI TY The presen t distrib utional p atterns of butter fly specie s divers ity in Ca lifornia are summariz ed in Figure 5- 3. Approximate numbers of endemic species and subspecies found in each area ar e indicated i n parenthesis. T his map is de rived from th e species di stribut ions described in detail in Chapter 8 of the present work. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 133 It is clea r from exa mination o f Figure 5 - 3 with Figure 5 - 1 that the overal l patte rn of butterfly diversi ty appro ximately p arallels the patte rn of plan t diversity in Califo rnia. The highes t diversity o f butterfl ies is assoc iated w ith the greates t topograp hic r elief, in the Sierra Nev ada Range of northern and central Calif ornia and the Transverse Mountain Ranges of southe rn California. Th e second highest dive rsity is located along th e coast, asso ciated with the various Coast Ranges of the state. A third c enter of b iodivers ity is in the north ern part o f the G reat Centr al Valley of C alifornia, w ell stud ied by Arth ur Shapiro a nd his st udents at the University o f Californ ia at Davis, and Sterling O. Mattoon and a notable group of co - workers, over the years. T he fourth center of butterfly biodiv ersity in th e stat e are the California d eserts. Intere stingly, th e Colorad o Desert area has a gre ater diversity th an the Mojave Deser t area, perhaps d ue to the altitud inal restrictio n and colde r winters of the north ern desert region. Nonetheless, it is perhaps somewha t surprising that the Great Basin desert regions of Inyo County rank last in biodiversity. [ The above brief discussion is subject to ex tensive modific ation after discussion with JFE and SOM, and c onstruction of Fi gure 5 - 3 on actual data figures, rather than impressions.] One area of California not included in the above discussion is the Channel I slands, lying offshore in the Pacif ic. Their butterf ly faunas are sufficien tly interesting th at we devote a separate short chap ter to them (C hapter 6 ). CONCLUS IONS The precedin g discussion paints o nly an overall picture of the p atterns of species d iversity in the sta te of Cali fornia, and forma l intensiv e analysis will occup y future g eneration s of lepidopterists for many y ears. In one sense, how ever, it is a race be tween the developer elements in the state and the natural ists and conservation ists to assemb le those data before the species ar e exterminated or extirpated in the mor e vulnerable regions. Fortunately, ge nerations of lepidopterists w ho went before us co llected acros s the state and even in early urban a reas, such as San Francisco, Los Angeles, and San Diego. By recording their captures in publications and in the form of specimens preserved in public museums today, we can reconstruct the biodiversity present a century ago. Without that huge data base to draw upon, we would have little knowledge of the remarkable species diversity that was once found in t he Los Angeles Basin and the San Francisco Bay area. One does not have to travel to the tropics to see dr amatic examples of environmental change and biodiversity loss; t hey are present right here at home in our own state. T he butterflies of California are an eloquent symbol, in many cases, of man's role in changing the natural environment. The great loss to mankind of once - wild places to ill - c onsidered development schemes and environmental degradation is pa infully evident even in the loss of thi s group of colorful w ildlife. It is incumben t upon us to not only record what we see today (for who can predict what tomor row will b ring in many o f these area s?), but also to work to mitigat e the effects of human ac tivities on L epido ptera popu lations in C alifornia. This las t topic is so importa nt that w e will cove r it in a separate chapter, Chapter 7, w hen we review extinct and threatened species found in the California butterfly fauna, conservation issues, and prospects for the future. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 134 Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 141 The Gulf F ritillary ( Agr aulis vanillae ) feeds on intr oduced Passion Flower vines that are grown as ornamentals o n trellises and backyard fences in urban areas. When freezes kill these vines back during the winter, the butterfly disappears, only to be reintroduced from Arizona or Mexico the fo llowing summer via migrants. Some butterflies have become more abundant in urban areas due to planting of thei r host shade trees. For example, the Mourning Clo ak ( Nymphalis antiopa ) has become relatively abundant in cities due to widespread planting of Weeping Willows, Poplars, and Ch inese Elm. The Western Tig er Swallowtail ( Papilio rutulus ) occur s wherever Sycamores and Modesto Ash have b een planted a s shade trees in parks and along city st reets. The Dusky- Wing Skipper ( Erynnis tristis ) has followed its host, C ork Oak, into suburban areas while a Crescent - s pot Nymphalid, Phyciodes phaon , ha s followed its host Lippia wherever Lippia lawns have been planted in u rban areas in the deser t, particularly a t Brawley and Bl ythe, but it has also been successfully introduced to Lippia lawns in H emet in the western pa rt of Rivers ide Co unty. Along with the spread of the Papi lio rutulus , due to the plantin g of Sycamore trees in parks, the arrival of paras itic mistletoe on t hese trees has made pos sible the wider estab lishment of the Great Blue Hairstreak, Atlide s halesus, whose larvae feed o n these mistletoes. Finally, it ought to be noted that not all butterflies have followed plantings of their host plants into urban areas. Canna Lilies are widely used as pl antings throughout southern California, yet the Canna Skipper, which has been repo rted several t imes as strays in so uthern California, has no t become establish ed to date. 2. Agricult uralization Like urbanization, the spread of a griculture normally results in the total loss of native habitat. In rare instances, narrow hedgerows of native vegetation are left between fields but normally only by farmers who are seeking to pre serve some cover for gam e birds, an ac t of wildlife conservation rar ely encountered in California. M ost native sp ecies, of cou rse, will disappear w hen their habitat and foodplants are removed. How ever, agricultural practices have introduced some species which serve as ex cellen t hosts for immense populations of certain species. The widespread planting of alfalfa througho ut the year in th e Imperia l Valley a nd Coachell a Valley of southeas tern California has made possible tremendous populations throughout the year of the Alf alfa S ulphur, Colias eurytheme . In addition, certain lycaenids such as Everes comyntas , Hemiargus gyas , and possibly Ministrymon leda have been more common due to the widespread plantin g of beans (genus Phaseolus ) as an ag ricultural crop in sou thern Cali fornia. Members of the cabbag e family have been planted fairly wide ly and Pieris species such as rapae and prot odice may become abundant at times in such areas. This family in cludes commerci ally grow n mustard (g rown for mus tard seed a nd creatin g musta rd spread). Fields of co tton (family Malvace ae) provide an alternat ive host for V anessa cardui , the Painted Lady, and the Gray Hai rstreak, Strymon melinus in both the so utheaste rn Califo rnia area and the Grea t Central V alley o f Califo rnia. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 142 In citrus orchards, two s pecies of swallowtails may become a pest at time s and have spread throughou t new agricul tural a reas as a resul t of their ab ility to sw itch to cit rus from oth er native foodplan ts. These in clude th e Anise Sw allowtail ( P apilio z elicaon ) and the G iant Swallo wtail ( Papilio cresphontes ), the la tter bei ng an invad er from Ar izona w hich origin ally ap peared in the Imperial Valley along the Colorado River and then moved up int o the Bakersfield area and north in the Central Valley. Extensive plantings of Sorghum grass and corn allowed the population sizes of the skipper ( Lerodea eufa la ) to increase, although they have not become of pest proportions. Irrigatio n canals in so me of the ma jor agri cultural are as have provided a d isturbed hab itat for cert ain butterfly foodp lants. Sun flowers (genus He lianthus ) do well in such habitats, and the Patch Butte rfly ( Chlosyne lacin ia ) multiplies abundantly in such favored areas. Likewise, agricultu re in th e Imperi al Valley ha s favored the growth of Alka li Mallo w ( Sida heteranea ??) along th e margins of field s. This is the hos t for seve ral skip per species ( Pyrgus scriptura an d P. communis ) as well as a hairs treak ( Strymon colu mella ), and these three butterflies h ave become relatively abundant in this field margin or hedgerow habitat. 3. Water Projects Water proje cts of all typ es have ef fected but terfly pop ulations in th e state, from d ams and reservoirs to harbors and bre akwaters , as well as wells tha t lower th e water ta ble, effe cting na tural surface water em issions. Dams and reservoirs no rmally create a total loss of habitat d ue to its submergence under water. The classic exa mple perhaps is Hetch Hetchy D am on the Tuolumne River in the Hetch Hetchy Valley just nor th of Yosemite Valley, inside Yosemite Nationa l Park. Short ly after the turn of the century, the city of San Francisco managed to pursuade the federal government to allow them to dam the Tuolum ne River he re insid e a nationa l park and flo od this g reat valley (larger than the present Yosemite V alley which is packed with millions of visitors every year). A hos t of butterfly h abita ts as well as a g reat numb er of other wil dlife sp ecies wer e lost by thi s flooding to create a reservoir for a water supply f or San Francisco. In the western part of the state, c loser to the coast, several di version and drainage pro jects have effec ted butterfly populat ions. There w as originally a system of na tural marsh es in the San Juaquin Valley, which probably had extensive colonies of t he ski pper Ochlodes y uma and its foodplant, the grass Phragmites communis . Today these colonies of the gr ass host and the butterfly have been virtually eliminated in the San Juaquin Valley by widespread drainage projects. Along the Kern River, at Weldon, the skipper butterfly Pseudocopaeodes eunus , has been greatly effected by water diversion projects above and below its colonies. These diversion projects effect the species adversely in some years by drying out the habitat, or p ositively in other years by protecting the habita t against floods . In the Sacramento River Valley, th ere are many species like this skipper, Ochlodes yuma . We have no idea y et what the impact will ultimately b e at periphe ral canals and the planned d iversion of the Feath er River water t o southern California. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 143 Another co mmon type of water pro ject in Califo rnia is deep w ells to tap undergro und water supplies. The effect of such wells i s to lower the water table as greater and greater amounts of water are withdrawn for human use. Along the San Jacinto River in the San Jacinto Valley, cottonwood trees are being lost due t o the dropping of the water table. On these co ttonwoods grow mistletoes which are the host plants for the Great Blue Hairstr eak, Atlides halesus . The d ramatic dropping of the water t able in the Owens Valley on the easte rn side of the Sierra has resulted in the drying up of springs there. This ha s definitely resulted in the extinction of the satyrid, Cercyonis pegala wheeleri , at Owens Lake, and possibly the extinction of colonies of the beautiful fritilla ry, Speyeria nokomis apacheana , south of the current known colonies in Round Valley. The dropping of the water table in deser t areas results in the drying up of isolated springs. Thus pupfish populations in Death Valley and elsewhere, especially in Nevada but also in California, have been threatened b y the loss o f water. So me butterfly species in t hese areas, dep endent on the ir host plants which in turn are dependent on a constant water supply in these desert springs, will die out like the pupfishes, for example the Phragmites grass and the Ochlodes yuma skipper are highly interdependent on each other in these iso lated desert cany ons where springs have occurred for thousands of years. Flood control channels built to control occasional flooding by major rivers such as the Los Angeles River in Los Angeles County and the Santa Ana River in San Bernardino County has resulted in extensive loss of riparian habit at and woodland, including willows and other foodplants of Limenitis lorquini , Satyrium sylvinus , and Phyciodes campe stris . Cloud seedi ng efforts over the Sierra Nevada s and attemp ts to increase sn ow pack may effect butterfly populations by influencing the increased ab undance or decline of their respective foodplants. Agricultural activities may effect weather too, by increas ing humidi ty in the Imperial Valley o f southeas tern Cali fornia. O riginally a desert, this ex tensively ag ricultur alized area may have an in creasing frequen cy of summe r thundersto rms due to the increas ed humid ity from evapora tion of irrig ation w ater, as w ell as trans piration from the g reatly in creased p lant bioma ss of these areas. These a gricultural and u rbanized areas now act as co rridors for nectar sou rces, water, and host pl ants fo r larvae for Mexican species to come up into s outhern C alifornia fro m eith er Ba ja Californ ia or Arizon a. Clearing th e vegetat ion along the M ojave River in the desert has destroyed Atriplex foodplants and thus endangered populations of the very rare Desert Blue , Plebejus emigdiones . Along the coast, the construction of breakwaters has stopped the natural creation of sand dunes or has resulted in an increase of dune building, depending on the locat ion of the breakwater. The construction of breakwaters in harbors then may reduce erosi on and loss of coastal sand in one area, bu t by changing coas tal currents may cause actua l loss in an other area. The fragile coastal sand dune habitat which supports various rare and high ly restricted subspecies of a number of blues, metalmark s, and fri tillarie s, can be greatly effecte d by man's activity o ffshore in an attempt to preserve beaches that have provided sheltered anchorages for recreational boats. 4. Road Construc tion Road con struction re sults in a compl ete loss o f habitat in the imm ediate a rea that th e road is occupying, of course, but the new roadside environment and roadcuts through hills provide a Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 144 disturbed habitat where secondary succession can occur. Richard Arnold (1983) notes that two Lupinus hosts for Ple bejus icario ides missionensis read ily colonize roadcu ts within a yea r or so of road construction and rapidly provide a habitat for expans ion of the butterfly's range. John Downey (1962?) cites this phenomenon for all Lupine hosts of the western icario ides subspecies. The natura l host bio mass for Papilio bairdii in the San Bernardino Mountains has been greatly favored by roadcuts where the host sage, Artemis ia dracunculus , is widespread in such habitats. A widely used species of wild buck wheat ( Eriogonum fasciculatum ) utilized by Apodemia mormo metalmarks and a number of blues is a frequent colon izer of roadcut s, such as on the p asses leading fro m the San Francis co Bay area to the Great C entral Valley o r on Cajon Pass f rom the San Berna rdino Val ley north into the Mojave Desert. H ere Eriogonum fascicula tum has foll owe d the new highways (such as Interstate Highway 1 5) out into the desert fro m the mountains and the San Bernardino Valley. An interesting ques tion for lep idopteris ts to inv estigate in the future w ill be whether butterfly species as sociat ed with this foodplan t will follo w it ou t into the d esert. Other common plants that incr ease in roadside habitats are t he mustard Brassica , which enhances the distribution of Pieris rapae and P. protodi ce , and various mallows ( Sidalcea in the family Malvceae) which serve as a host for various Pyrgus species (including P. he liopetes ) which follow its distribution. According to Richard Arnold (1983), the legume L otus formosissimus is doing well along the roadsides near the only know n colony of the Lotus Blue ( Lycaeides idas l otis ) on the Mendocino County coast. The fact that the host plant is spreading may expand the range of this endangered butterfly due to the presence of r oadcuts near the sole known remaining colony. A much more co mmon bu tterfly, the Anise Swallow tail ( Pa pilio zelicaon ), has followed the expansion of wild fennel ( Foneniculum vulgare ) which does well on roadcuts such as in the San Francisco Bay area. Roads, p articular ly in fore sted areas , act as na tural co rridors for butterfly flight p atterns , much as do canyon bottoms and stream s. The existence of a road may also i ncrease runoff of rainwater from the highway along the roadsides, allowing for less vegetation growth there. Both larval host plants and nectar sources may become quite abundant. In the higher mou ntain areas, where snow plows pile up snow along the roadsides duri ng the winter, the increased snow pack there provides greatly enhanced precipitation for spring and summer plant growth. Even in the desert, plant growth along desert highways is noticeably improved, including such plants as Sphaeraldea (fa mily Malvaceae) wh ich serves as a host for the skipper, Pyrgus scriptura . In northern Ca lifornia Sedu m spathelfolium does wel l along roadsides and the hairstreak, Incisalia mossii (various sub species in different ar eas), may coloniz e these abundant hosts. L ikewise, buckwheats or Eriogonum species in general ( E. nudum , parvifolium , umbellatum , etc.) tend to do well on roadcuts and promote the establishment of new colonies of blues and metalmarks. Finally, the exist ence of the road increases coll ecting pressure on the co llecting local ity. A classic exa mple of this is the fact that the Tioga P ass Highwa y (right outs ide the eas tern ent rance to Yosemit e National Par k) makes it the mo st heavily co llec ted spo t for alpine species in the state of California . Fortunately, there h as been no apparent effect of this heavy collecting p ressure each June and July to date. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 145 Finally, two other examples of spec ific butterfly expansions due to roadcuts may be found in southern California mountains. In the San Bernar dino Mountains, the foodplant of the rare marble Euchloe hyantes andrewsi , which is S treptanthus bernardinas , follows roadcuts and provides an egg and larval substrate, wh ile at the same time the road acts as a natural corridor for the ovipositing marble females (Chris Henne, pers onal communication). Likewise, the legume Lotus scoparius does well on r oadcuts and provides the expanding resource for the hair streak, Callophrys perplexa , and the blue, Glaucopsyche lygdamus australis . 5. Grazing Grazing has a number of varied effect s on butterfly populations. First, it can cause chan ges in the grass species composition of the commun ity in the gra ssland area. The nativ e perennial bunchgrasses tend to be replaced by introduced a nnual European grasses. This possibly influences the species of skippers and Satyrids th at live in these grassland communities, although how much influence is exerted is still unstudied. A more d irect immed iate eff ect is that erosio n normally increas es with ove rgraz ing. Additionally, certain species of plan ts poisonous to cattle, such as Astragulus , Lupinus , or Asclepius milkweeds, may proliferate due to the impact of grazing. Butterflies that feed on these poisonous plants may actually become more abundant due to the grazing pressure. Species that are specializ ed to feed o n oak trees, however, may no t fair well in o ak grassland areas where overgrazing occurs. Oak seedlings are prevented from growing, and the oak trees will eventually die out in such an area. Natural meadows, particularly in mountain area s such as alpine meadows in the Sierra Nevada and Garner Valley in the San Jacinto Mountains of southern California, may become so overgrazed that they are invaded by weedy species, and today we have n o idea w hat the original flora and fauna was of these ric h, wet, boggy meadows, before they were overgr azed by cattle and sheep. Such grazing pressure was obse rved very early by John Muir in the 1880's and 1890's in the Sierras when he helped herd sheep before becoming an ardent conservationist. The divers ion of wa ter sources fo r cattle tr oughs an d springs fo r irrigatio n projects m ay cause meadow s and grasslands to dry up and essentially di sappear from thei r natural state. Grazing in marginal habitats, su ch as the eastern Mojave D esert, makes no sense at all. The diversion of natural water sources and the grazing of catt le in a desert environment today impacts heavily o n the native desert vegetation and resu lts in very few cattle being raised w hile creating inestimab le damage to the environment. The grazing of wild burros in th e western desert mountain ranges in higher desert are as has considerable impac t on grasses and brush, as well as grea tly disturbing natural springs. (J.F.E.: Look up Kingston Range and Granite Mountains Resource Su rvey Grazing Report, and report on eff ective mining on plants for T.C.E. to use or for J .F.E. to modify this section and the later one on mining.) It also should be noted that changes in the spotting pattern of the Satyrid butterflies, Cercyonis oetu s , have been noted in Colorado populations living in heavily grazed meadows. Thus, the evolutionary and ecological impact of grazing may effect genetic diversity and phenotypic charac teristics of butter flies as well as endanger their very co ntinued existence. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 146 6. Smog and Air Pol lution The effects of air poll ution have been reasonably well studied in plants where the deleterious effect on ponderosa p ine forests so me 60 miles east o f the Los Ang eles Basin have been noted. But the impact on butterfly populations has not really been studie d yet. It has b een hypothesized (Emmel and Emmel, 1973) that one factor possibly accounting for the decline in Phoebis sennae sulphur populations in the Los Angeles area has been the change in air quality. 7. Pesticides an d Herbicides The widespread use o f pesticides in agriculture, mosquito control, an d forest insect p est control is done virtually without a thought as to possible side effects on non - target insects and other wild life. Vast sp raying progra ms to control in jurious in festations or outbreaks may result i n incalculable effects on wild butterfly populations. To da te however, hard evidence from experimental studies is available only from one study (Emmel, 1986, 1988) in Fl orida where the effect of mo squito cont rol pestic ides on the Schaus Sw allowtail, a federally endangered species, and the common Giant Swallowtail ( Papilio cresphontes ) in the Florida Keys has been studied. Other chemicals of great concern are herbicides which are used along roadsides throughout the state and in the past, have been used on fir ebreaks to control vegetation growth. The direct impact can be measured in the loss of larv al and adult h ost and nectar pl ants, but a ligh ter impact may be a poisoning of the only populations of many of these butterfly s pecies in an area. As has been noted above, many species follow disturbed habitats along roadsides and as more exten sive native habitat is destroyed, roadside habitats may provide the last refuge for a number of these species. Periodic spraying by herbic ides to contro l roadside growth can only result in a t hreat to the continued existence of these spec ies. 8. Mining Mining has been rather extensive in many areas in California, starting in the days of the 1849 Gold Rush and continuing to date. O ne of the classic examples of ecological damage done by mining i s that of pla cer mining i n the Sierra n foothills. Placer min ing created hug e piles of gravel tailings which in some cases have remained barren of vegetation for a century and have changed the routes of watercourses as well. Lik ewise, modified strip mining, as in the Kingston Range of the east ern Mojave Desert (J. F.E. cite that study here.) has created wholesale habitat destructio n for the talc mine. Sand mining in the sandhills of southern Santa Cruz County [ask John Lane for details] is destroying unique ecotypes of the metalmark, Apo demia mormo, a nd the blue, Euphilotes enoptes , which feed on Eriogonum nudum in the sandhills. A population of the skipper Polites mardon on serpentine in Del Norte County is endangered by planned mining activities there [John: Get details from S.O.M.]. Quary mining on the eastern side of the San Bruno Mountains is destroying the habitat of the endangered hairstreak s pecies, Incisalia m ossii bayensis . 9. Ski Runs and Ski Ar eas Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 147 The construction of ski areas and ski runs through national forest s and other areas maintains a subclimax meadow or open slopes, preventing the regrowth of trees in t he cleared areas. There is probab ly a greate r grow th of wildf lowers in th ese areas than i n c limax forests, the reby increasing the number and variety of necta r sources and diversity of larval host spec ies of plants for butterflies. T hese areas also act as corrido rs for montane species su ch as Chlosyne hoffmanni , Papilio indra , Parnass ius sp., and Lycaena coppers. In particular, Lycaena cupreus , L. nivalis , and L. editha do well in these corridors created for skiing. Thus, the construction of ski areas commonly results in a greater abundance of butterflies during the summer months. 10. Forest Cutting and Lumbering Selective lumbering introduces second - growth vegetation which provides normally a greater abundance of nectar sources and larval foodplants for butterflies . Clear cutting, on the other hand, creates erosi on of slopes and siltation of streams w ith great ef fects on ripa rian habita ts downstream as well as the mountain slopes where the forest once stood. It would be particularly interesting to study redwood forests or o ther areas where clear cutting is taking place, in o rder to document what happens with species diversity among butterflies. For instance, in a redwood forest Pieris napi has a subspecies with very limited distribution in Humboldt County, along with the Anglewing butterfly, Polygonius faunus fulvescens . These butterflies are undoubtedly effected b y the logging operations in the forest, and it would be valuable to understand how short - term or long - term thes e effects a re. Lumbering also cre ates a direct ef fect on species th at use conifers as foodp lants. However, in every case in C alifornia, the b utterflies that f eed on conifers in th e larval stages a re widespread species an d lumberin g has only a temporary effec t in the im mediate area on their abundance. On the other hand, any fritillaries in t he genus Speyeria or other butte rfly species th at require a shade d forest cano py as a habit at for ov iposition activit ies, may b e extermin ated or g reatly reduced in numbers due to extensive lumbering in an area. The fact of the matter is that no detailed ecological studies of butterfly diversities have been done in lumbered versus unlumbered forests and such research is badly needed in not only our state but also the other western sta tes. 11. Fires The only study that we know of on the effect of fires on butterfly succession and diversity is that by Richard Kelton (1987, unpublished paper). Kelton studied fire in the chaparral area and listed all the species favored by fire along with their hosts. Fire opens up the habitat to allow both the hosts and the butterfl ies to increase. Fo r example, the southern California speci es Papilio indr a pergamus has a foodplant which does extr emely well on burned slopes. In the years following a forest f ire, this but terfly greatly expands in number due to the abundance of its foodplant. All the chaparral host specie s and their butterflies are prob ably benefited by fire because mos t of the plants are crayon - sprouters, and new growth would be favored by ovipositing females in any year, let alone following severe fire. The basic question in the case of severe f ires is whether there is any survival a t all of th e butterfl ies in th e area of a fire. Lycaenid a nd Papilio indr a pupae are normally placed in sheltered ar ea s and probably survive a fast moving fir e without much difficulty. Other species that pupate on dry foodplants, such as Pierids (Orange tips, Whites, etc.) probably have to reinvade fire -burned area from an adjacent unburned habitat. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 148 Besides chaparral fir es, we also have forest fires w hich cause wholesale destruction of an area, but then succession occurs on the burned slopes and, as in slopes cleared by lumb ering or for ski areas, butterfly host plants and nectar sources do quite wel l in the early years of succession. Grass fires in serpentine grassl and or other relativ ely open and sparse ly vegetated habi tats such as deserts will normally sweep through too rapidly to create long- term prob lems for b utterfl ies. In the New Y ork Mountains in the eastern M ojave Desert, the re is a very inter esting situation th at may be th e resul t of fire. Several plants , includ ing Purshia and Ceanothus greggii may represe nt relics from earli er burns in this mounta in range . Yet, there are no bu tterflie s associated with them (one would expect the hairstreaks, Satyrium behrii on the purshia, a nd Satyrium saepium on the Ceanothus species). On the co ast of Californi a, these butterfly speci es re- establish themselves on the newly sprouting Purshia and Ceanothus bushes quite readily. In the more isolated New York Mountains, a recolonizat ion may be a relatively rar e event for these hairstreak sp ecies. 12. Off- Road Vehicl es Recreational vehicl es for off - road trav el have had a co nsiderable impact on several habitats in Califo rnia. O ne of these ha bitats is the desert, where moto rcycle tra ils an d RV trails at Jawbone Canyon in the Mojave Desert and other desert mountain ranges (sites for Papilio indr a subspecies and other rare desert butterflies) have torn up the desert habitat extensively. At the Mari na Dunes in Monterey County and the Oso Flaco Dunes in San Luis Obispo County along the coast, motor vehicles racin g cross the dunes have caused the direc t destruction of plan ts which are t he hosts of an ecotype of Euphylotes enoptes tildeni there at th e Marina Dunes on Eriogonum latifo lium , and a subspecies of the metalmark A podemia mormo on the same plant. At the Marina Dunes exist s the only known sympatric populat ions of the two green hairstreaks, Callophrys dumetorum and C. perplexa . These have long confused California lepidopterists and taxonomists, and have only recently bee n differentiated as distinct spe cies by careful ex amination of the ir life h istories. It would be a tragedy to lose the one natural laboratory where the two species come in contact, and be unable to study possible hybridization and barriers to interbreeding that may exist between these beautiful butterfli es. At Oso Flaco Dunes, a beautiful ch eckerspot subspec ies of Thessalia l eanira is endangered by off - road veh icle activ ity. The rare blue P lebejus icarioides moroensis which inhabits the sand dunes in San Luis Obispo County along the coast, have been mostly destroyed by development and it would seem particul arly ironic if the last known populations were destroyed by man's careless us e of off -road vehicles in the sand dunes. 13. Introduc tion of Exo tic and Wee dy Plant Sp ecies into N ative Habit ats Under this category of human influence, we mention the alteration of vegetation by the accidenta l or del iberate i ntroductio n of plan ts from ou tside th e state o f Californ ia. In so me cases, these plants have increased butterfly abundance such as fennel ( Fo en icul um vulgare ) , which has made possible the increased distribution and abundance of the beautiful Anise Swallowtail. Likewise, th e Malva species fro m Europe have beco me widely distrib uted in distu rbed areas, roadsides, vacant lots, and agricultural are as greatly effec ting the dis tribution of Vanessa cardui and Vanessa carye . The distribution of mustards in the genus Brassica, especially European Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 149 mustards, has greatly increase d the n umbers of the white s in th e genus P ieris, esp ecially P. rapae and P. protodice . On the negat ive side of th e ledger, Tamar isk has been in troduced into the d esert areas w here it invades desert springs and riparian habitats and replaces cottonwoods, willows, and mesquite ( Prosopis ), all foodplants of native butterflies. G orse is becoming very abundant in northern California, like Scotch Broom has invaded much of the montane areas of southern California along roadsides. In the B ig Sur area, pampas grass has become naturalized and is crowding out virtually everything else. Ice plant on the El Segundo Sand Dunes, as well as a native but introduced bucwheat species, Eriogonum fasciculatum , have been crow ding out the native hosts of the rare El Segundo Blue ( Euphilotes battoides allyni ). Currently, an active removal program has been initiated in the El Segundo Dunes in order to remove these exotic species. Even in the remote desert areas, butterfly dist ributions can b e affected by accid ental or deliberate introductions. A colony of watercress was introduced some time ago at Paphalka Spring on Clark Mountain in the eastern Mojave Desert, apparently with some eggs of Pieris rapae on it. Today, a thriving colony of Pieris rapae occu rs in this most unlike ly desert habitat. L ikewise, th e widespread distribution of the vine of the morning glory, Convolvulus arvensus , may u ltim ately lead to Phyciodes picta becom ing establis hed in California . James Scott has found th at it fe eds on this Convolvulus in addition to its nativ e host. The butterfly is common in Arizona an d will undoubtedly eventually invade the Blythe area along the Colo rado River and find its Convolvulus foodplan t awaiting it for succes sful introd uction o f the butte rfly. Raven (1977) has provided a fasci nating summary of the literature on the introduction of exotic plants into California. This b egan in May 1769, when Father Junipero Serra r eached San Diego Bay and founded the first permanent European settlement i n what would be ultimately the state of C alifornia. At l east 16 spec ies of exotic plants were established in Califo rnia during th e Spanish colonization period of 1769 to 1824. During the Mexican occupation, from 1825 to 1848, 63 additional species were introduced. Some 55 more introduced plants arri ved during American pioneer settlement from 1849 through 1860. Thus by 1860, at least 134 alien species of plant s were established. By 1920, 186 immigrant species excluding odd fugitive plants from gardens a nd waifs from w ind- blown seeds, in southern California alone. In 1925, Jepson believed that 292 species of established exotics were in the California flora. By 1951, 437 introduced species of weeds were listed for the state, and Munz and Keck (1959) l isted 797 species of introduced plants. By 1968, Munz added 178 more species of introduced plants, for a total of 975 intr oduced species in California. Many new w eeds are being established in California w ith each passing decade, but not all are b ecoming a part o f t he naturalized flor a of the state. The good climate and widespread irrigation du ring the dry sum mers allows many species to p ersist and ev en reproduce by seed locally but this is due in part to human interference. Rave n (1977) concluded that there were a ctually 654 species of introduced plants in C alifornia at that time, ignoring those that were persisting with human assistance. Of these, 544 were of Old Wor ld origin and only 110 were of New World origin, the la tter includ ing some 5 0 from the Unit ed States, 23 from tropic al America , and 37 from South America. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 150 Chapter 8 SPECIES ACC OUNTS OF T HE BUTTERFLIES A ND SKIPPERS OF CA LIFORNIA P A P I L I O N I D A E The family Papilionidae, or swallowtails and parnassians, contain s three recognized subfamilies. The subfamily Papilioninae includes the true swallowtails, with about 517 species worldwide. The subfamily Parnassiinae contains the parnassians, which are found largely in the Holarctic and include 54 recognized species. The third subfamily, Baroniinae, contains only a single species ( Baronia brevicornis ) which is endemic to central Mexico (Heppner, 1991). The family is separated from its closest relatives, the Pieridae, and the other families of butterflies by having (1) six functional legs in both sexes; (2) a spur called an epiphysis on the tibia of the front (or prothoracic) legs; (3) a larval osmeteriu m or forked organ that protrudes forward from behind the head when the larva is disturbed; (4) five - segmented feet, with the tarsal claws single and simple in structure; (5) clubbed antennae which are not curved at the tip; (6) a head as wide as the thorax , and bearing short palpi; (7) a central (or discoidal) wing cell between the veins in both the forewings and hindwings which is completely closed (i.e., the veins are joined) at the cell end closest to the body; (8) larvae which are generally cylindrical, although often with an enlarged thoracic region in the younger stages of the true swallowtails. Most of the species of this family in California bear tails, excepting only Battus polydamus an d the two species of Parnassius . * * * * * * * * * Subfamily Papilioninae Latreille l. Papilio bairdii W. H. Edwards ADULT B EHAVIOR : The males of this wester n black swallowtail are s trong hilltoppers wh ile females are fre quent ly enco untere d flyi ng in ca nyon bottom s in se arch of t he foo dplant. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 253 ocell us is la rger t han the a nte rior oce llus o r is e qual in s ize , wherea s in sthenele the posterior ocellu s tends to be smaller than the anter ior ocellus. GEOGR APHIC AN D There i s cons iderabl e ge ographi c vari ation i n Cercyonis sthene le , as SEASONA L VARIAT ION: exemplified by the fact t hat there are seven subspecies represented in the state of Calif ornia. Genera lly, the pattern of v ariation is more straightforward in this species than in the preceding one (see C. pegala ), prim arily be cause popul ations tend to be mo re wide sprea d and are not highl y is olate d by bei ng conf ine d like pegal a to discre te boggy meadows or other limited habitats. The re is no seasonal variation because the species is univoltine and has a relatively short emergence period thro ughout i ts ra nge i n Cal ifornia . GENERAL REM ARKS: As with the other Cercyonis , co nsidera ble f ield w ork re mains to be done on the ecolo gical a nd evol utiona ry fa ctors t hat co ntrol a nd co ntribu te to t he rema rkabl e variat ion in ocel lation a nd othe r wing c hara cters in the adul ts. The native grass species used by C. sthenel e and the other Cercyonis species remain to be determined. This Cercyonis is one of our most successful s atyrid butterflies and occup ies a variety of habitats across California. It is character istically a lowland s pecies, flyin g in oa k grass land or hot se mi - desert areas. Considerable geographic variation occurs. I t also has th e distinction of h aving been the first Californ ia butterfly (in the form of the nominotypical sub species) to be driven to extinction by di rect hum an act ion. a) C. s. sthenele (Boisduval ) DIAGNOS IS : The nominotypical subspecies was characterized by a heavily marked median band o n the unders ide of the ventr al hin dwings , wit h exte nsive white scal ing on the dis tal half of the wing.. ORIGINA L DESCR IPTION: Boisduv al, Ann. Soc. Ent . France , (2)10 : 308 ( 1852). TYPE L OCALITY: San Fra ncisc o, Ca liforni a. Le ctotype in U.S . Nat ional M useum, desi gnate d by F. M. Brow n, Jou rnal of Rese arch on the Le pidopt era , 4: 141 (1966 ). DISTRIB UTION : The n ominot ypica l subs peci es was found only i n the area now oc cupi ed by t he city of San Francisco and has been extinct since approximately 1880. There are very f ew spe cimens survi ving in colle ctio ns, most havi ng bee n in the Cali fornia Academy of Sciences collec tion and thus destroyed in the fir e that followed the San Fra ncisco earth quake of 1906. ELEVAT ION RANGE: Sea l evel t o perha ps 5 0' eleva tion. HABITA T AND PLANT COMMUN ITIES: Unknow n, but l ikely gra ssy slope s wit hin Nort hern Oa k Woodla nd. PHENOL OGY : The onl y detail s of C. s. sthe nele 's biolo gy that were ever recorde d were t hat it flew i n the month of Ju ne, ha d onl y one gener ation a nd p refer red to "sett le on t he groun d" rathe r than " on the unde rside of oak bra nche s" like silvestris . It is appare nt tha t sthenele pas sed int o oblivi on due to man 's dest ructi on of its restricted habitat on the San Francisco peninsula. FOODP LANTS: U nknown. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 254 b) C. s. silvestri s (W. H. Edwards) DIAGNOS IS : This tax on is represe nted b y a se ries of weste rn slo pe Sie rran po pulat ions tha t share a gene ral l ight br own gr ound c olor a nd sma ll num ber o f ocell i on t he v entral surfac es of t he w ings. A mar ked me dian ba nd is lacki ng. ORIGIN AL DESC RIPTIO N : W . H. Edwa rds, Proc. Ac ad. Nat. Sc i. P hiladel phia , 13: 162 (1861). This subsp ecies was misp laced by Miller a nd Brown (1981 ) under Cercyo nis oetus . TYPE L OCALITY: "Cal ifornia ." Lect otype i n Carnegi e Mus eum, desi gnat ed by F. M. Brown, Trans action s Americ an Ent omolo gical Societ y , 90: 36 2 (19 64). Type local ity specifically fixed by Emmel, Emmel and Mattoon, Systematics of Western No rth American Butterflies , : (1998) to lower Feathe r River Canyon, P lumas County, California. The types were collected by Lorquin, given to Behr, and transmitted to Edwa rds, who de scribe d it. DISTRIB UTION : This s ubspeci es is found o n the west sl ope of t he Sie rra Ne vada fr om no rthern Kern Coun ty north to the Mount L assen region . To the south, it blends with subspecies beh rii ; to the north, it blend s with subspecies sineoc ellata . ELEVAT ION RANGE: From a pproxi mate ly 1,5 00 to 5,000' e levat ion, m ostly below 4,00 0'. HABITA T AND PLANT This subsp ecies is typically foun d in grassy areas with in Foothill COMMUN ITIES: Woodla nd, Mi xed C hapa rral, o r lo wer ed ges o f the L ower Monta ne Conife rous F orest . PHENOL OGY : Univolt ine, with di apause in the ne wly hat ched fi rst - i nstar lar vae. T he adult s begin e mergi ng i n early June and t he fli ght c ontinue s th rough m id - Aug ust, wit h a peak i n earl y Jul y. FOODP LANTS: Pr esumably grasses, as yet undetermined. c) C. s. behrii (F. Grinnell) DIAGNOS IS : The status of the subspecies behrii has been unclear for many y ears because of the dest ructi on of t he ori gina l type se ries a nd lac k of top otypi cal mat eria l until recent ly. Em mel, Em mel, and Mat toon ( 1989) have de signate d a ne otype of behrii an d have sho wn tha t thi s is t he na me whi ch re fers t o pop ulati ons of sthenele in the California Coast Ranges. There are characterized by a dark basal half of the ve ntral hindwi ng, wi th a c ontrast ingly l ighte r di stal ha lf of t he hi ndwing , and a lack of ocelli. ORIGINA L DESCR IPTION: F. Grinnell , Entomologica l News , 16: 33 (1905). TYPE L OCALITY: Mount Tamal pais, Mari n Count y, Cal iforni a. The ori ginal type was lost in t he 1906 Sa n Fra ncisco eart hquake and fi re. Ne otype desi gnated from t he type locality by Emmel and Emmel, Systematics of Western North American Butterflies , : (1998) a nd dep osited i n the Natura l His tory M useum of Los Angele s Cou nty. DISTRIB UTION : This s ubspeci es is found i n the C alifor nia Coa st Ran ges fr om San Diego County north ward t o appr oximate ly Hu mboldt Cou nty. N orth of Humb oldt C ounty, it blends into the s ubspec ies sinoecellata . To the e ast i n the s outhe rn port ion of its range (e.g. , the Tehac hapi Mount ains), it bl ends with t he we st - slope Sierran subspecies silvestris . Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 255 ELEVAT ION RANGE: From nearl y sea l evel to appr oximate ly 5, 000' el evatio n. HABITA T AND PLANT The typi cal habi tat i s woode d slope s or ca nyon bottoms withi n the COMMUN ITIES: Northe rn or Southe rn Oak Woo dland com munit y, near ed ges of Mixe d Chapa rral, a nd ope n are as nea r Nort hern a nd S outhe rn Mixe d Evergreen Forest. PHENOL OGY : The flight period is from early to mid - June into early Augus t, with a peak in early July. FOODP LANTS: U nknown. d) C. s. paulus (W. H. Edw ards) DIAGNOS IS : A well - ocellated subspecies w ith silvery gray mottling on th e underside of th e wings. It occurs in the western Great Basin areas just inside Calif ornia. ORIGINA L DESCR IPTION: W. H. Edward s, Canadi an Ent omol ogist , 11: 50 (1879). TYPE L OCALITY: "Nevada" , rest ricted to Virginia C ity, Sto rey Count y, Nevada b y F. M. Brown, Trans action s Ameri can E ntomol ogical Soc iety , 90: 365 (1964). Neotyp e in Carne gie Muse um, de sig nated b y F. M. Brown (196 4). DISTRIB UTION : In Californ ia, the subspecies paulus is restricted to an area east of t he Sierra Nevada crest , from Mono C ounty s outh in to the n orther n half o f Inyo County , mostly along the ea st slo pes of t he Sie rra N evada b ut wit h popul ations e xte nding into the Wh ite Mountains as well. T he exact eastern a nd northern lim its of its range are unclear at the present time. Populati ons from areas east and southeast of paulus ' range show some blending towards the facies of the typical western Colora do su bspeci es mas oni (Cross). ELEVAT ION RANGE: From 4, 500 to 7, 000' ele vatio n. HABITA T AND PLANT This s pecies is t ypica lly f ound i n the Great Basi n Sage brus h Scr ub or COMMUN ITIES: Pinyon - Juniper Woodland plant commun ities. Its favored hab itat seems to be small dry canyons and washes, particularly where nectar sources such as Melilotus (sweet clover) occur. PHENOL OGY : Recor ds exte nd fr om la te Ju ne int o ear ly Au gust, with a peak flig ht in mid - July. The species overwinters as a fi rst - instar larv a. FOODP LANTS: U nknown. e) C. s. masoni (Cross) DIAGNOS IS : A subspecies of sthenele with a striated, u mber - brown unde rside beari ng a strong median band a nd pro mine nt ocel li. Fr eshl y caught male s ofte n have a flush of rusty c olor on the under side of the f orewi ng at t he ape x. T his red dish t int soo n fades after death. ORIGINA L DESCR IPTION: Cross, Pr ocee dings C olora do Mus eum N atural Hist ory , 16(1): 9 - 10 (1937). TYPE L OCALITY: Blac k Ridge B reaks , Mesa Count y, C olorad o. Holot ype i n Denve r Muse um of Natural History. Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 256 DISTRIB UTION : This wi dely dist ribute d subs peci es is fo und from t he ca nyon c ountry of e xtreme wester n Color ado west ward thr ough Uta h and southe rn Neva da to sou theast ern Cali fornia an d also into nort hern Ari zona and e xtreme n orthwest ern New Me xico. Within Californ ia, it is fou nd in t he hig her mou ntains of the e aste rn Moja ve Desert , such as the Granit e Mounta ins nort h of Ambo y, the Pr ovide nce Mount ains, the New Yor k Mounta ins, the Clark M ountai ns, and t he Kingst on Range. It is also presen t, at least in a blend - zone, wit h sub species pau lus in the Inyo Mo untai ns, Last C hance Range, and Gra pevine M ounta ins of Iny o County . ELEVAT ION RANGE: From a pproxi mate ly 4,0 00' t o 8,000' eleva tion. HABITA T AND PLANT This subsp ecies is typically found within th e Pinyon - Junipe r Woodla nd COMMUN ITIES: of the desert mount ains, both on ope n slopes and i n can yon bot toms. PHENOL OGY : Our recor ds extend fr om mid - June to early August, with a peak in early July. FOODP LANTS: U nknown. f) C. s. sineocellata Aust in and Emmel DIAGNOS IS : A subsp ecies of extreme northern California in the northwestern Gre at Basin, characterized by a whitish to light grayish under side that generally lack s ocelli. ORIGINA L DESCR IPTION: Austi n and E mmel, Systematics o f Western North American Butterflies , : (1998). TYPE L OCALITY: West si de of C rump La ke, 8. 2 mi. N. A del, L ake Count y, Ore gon. H olotype in Nevada State Museum. DISTRIB UTION : This subspecies is r ather widely distributed in the Great Basin areas of north wester n Neva da and n orthea stern C ali fornia, n orth ward int o sout heast ern Oreg on and westwa rd to t he Kl amat h Mount ains a ssembl age a nd t he sout hern Casca de Ra nge of O regon. ELEVAT ION RANGE: From 2, 500' to 6, 000' ele vati on. HABITA T AND PLANT Within th e Great Basin area , most populatio ns are found with in the COMMUN ITIES: Great Ba sin Sa gebr ush Sc rub or P inyon - Juni per Woodl and communities. Westward in the Klamath Mountains and southw estern Oreg on popul ati ons, the h abita t is typi call y Northe rn Oak Wood land or open are as withi n Nort hern Mixe d Everg reen Fo rest. PHENOL OGY : Adult captures are recorded from mid - June to early August, with a peak in early to mid - July. FOODP LANTS: U nknown. g) C. s. hypoleuca Haw ks and Remington DIAGNOS IS : A maritime su bspecies of Santa Cruz Island of f the coast of California, superficially r esembling th e extinct nomino typical subspecies sthenele with a strongl y acce ntuat ed ve ntral median band; specime ns ave rage la rger t han nominotypical sth enele . Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 257 ORIGIN AL DESC RIPTIO N : Hawks and Remi ngton, Systematics of Western North American Butter flies , : (1998). TYPE L OCALITY: , Santa Cruz Island, Santa Barbara County, Californ ia. Holotype in Natural Histor y Muse um o f Los A ngel es Cou nty. DISTRIB UTION : This s ubspec ies ap pears t o be c onfine d to Sa nta C ruz Isl and, altho ugh t he possi bilit y of a p opulat ion on Santa R osa Island re mains to be invest igate d. ELEVAT ION RANGE: From sea level to approxim ately 2,000' elevation. HABITA T AND PLANT The sub species is found within Southern O ak Woodland, on slop es and COMMUN ITIES: in canyo n bottoms. PHENOL OGY : Capture s are re corde d form early J une to e arly Jul y, wi th an a pparent peak fl ight in late June. FOODP LANTS: U nknown. 50. Cercyonis oetus (Boi sduval) ADULT B EHAVIOR: This small species flies w ith a weakly bobbing f light, frequently stopping at yellow composite flo wers such as Sen ecio and ra bbitbr ush ( Chry soth amnus ) in the grassy sagebrush areas. Adults mate for approximately one hour and the female s lay th eir egg s singl y, depos itin g about 10 to 12 e ggs a da y duri ng thei r lifetime of as much as four weeks. Th is species frequently settles on b are gravelly slopes , w here its mottled gray unders ide camouflages the butterfly nicely. LIFE H ISTORY: Th e cr eam - colored egg is bar rel shape d, wit h about 28 prono unced vertic al ribs and ver y fine horizo ntal c onnect ing ribs . The ma ture l arva is simil ar to t hat of Cercyonis sthenele , except that th e dark green mid - dorsal line is bordere d by a thin white line on each side. The pupa is similar to those of the other Cercyonis specie s in C ali fornia, and s hows va riat ion in color rangi ng fr om light gree n wit h no striations to d ark brown with heavy linear marking s. CHROM OSOME NU MBER: n = 27 (T. C . Emmel , unpubli she d). SIMILAR S PECIES: See Cercyonis pe gala and C. sthene le for di scussi on. GEOGR APHIC AN D Most of t he s ubspeci es of Cercyonis oetus are generally weakly SEASONA L VARIAT ION: differentiated and intergrade. H owever, pallescens is quite d istinctive fro m most oetus , char on (W. H. E dwar ds) (R ocky M ountai ns), a nd phocus (W. H. Edwards) (Br itish Columbia and the extreme north wester n U.S.). Our single subspe cies on the West Coa st in the Sierra Nevad a, Cercyonis oetus oet us , grades towards pallescens in Modoc C ount y, as disc ussed i n the distr ibutio n sect ion a bove. The re is no sea sonal varia tion i n this univol tine s pecie s. GENERAL REM ARKS: Ext ensive mar k - recapt ure st udies ha ve be en do ne by T . C. E mmel ( unpub lishe d) in Colo rado populati ons, whi ch sh ow that the butt erfly c an pe rsist i n even ve ry small populat ions from yea r to yea r (som etime s fewer t han 50 in dividua ls represent a stable populat ion ). The se loc al p opulat ions m ay e xhibit consi derabl e variat ion ove r short distanc es in the spot ting pa ttern on the un derside of the wing. Some la rger populat ions ma y num ber tens of thousa nds of i ndivid uals an d sprea d out ove r many square mi les. These studies have also shown that females and males may live as three to fo ur weeks in the wild, though th e usual lifespan is Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon 258 probabl y sh orter. Sim ilar s tudies need to be done in C alifor nia p opulat ions by intere sted in divid uals. St udi es of nat ural pr edation on the Colora do oet us adults suggest s tha t bi rds and other preda tors may ha ve thei r att tac ks defl ecte d by t he spotting pattern on the hindwing. These spots apparently offer a tempting tar get for bi rds to grab at and t hus a llow the butt erfly t o esca pe, l eaving behind only a sm all triangle of broken wing section in the bird's beak. This small w estern Cercyonis species is occ asional ly co nfused with Cercyonis sthenele, but it can be immediately dis tinguished by the fact th at the posterior ocell us on the unders ide of t he fore wing i s clos er to the wing ma rgin t han the anterior ey espot (in C. st henel e , the two ventral fo rewing ocelli are equ ally distant from t he wi ng ma rgin). Als o, on the unders ide o f the hindwi ng, t he in ner ma rgin of the me dian ban d of oetus is strongly lob ate. a) C. o. oetus (Boisduval) DIAGNOS IS : The unders ide of oet us is gray to gray - brown, and the mesial band is well marked by dar k bor ders tha t are shar ply in dente d in a zig - zag pattern throughout their lengths. ORIGINA L DESCR IPTION: Boisduv al, Ann. Soc. Ent . Belgi que , 12: 63 (186 9). TYPE L OCALITY: "California. " Lectotype in the United States N ational Museum, des ignated by F. M. Brow n, Jou rnal of Rese arch on t he Le pidopt era , 4: 142 - 143 (196 6). Emmel , Emmel, and Mattoo n, S ystematics of Western North American Butterflies , : (1998 ) desi gnate d the t ype l oca lity as Hwy. 70 at Bet terto n Cre ek, 6. 5 roa d mil es W. of P ortol a, 500 0' ele vation , Plum as Cou nty, C alif ornia. DISTRIB UTION : In California, the species tends to be found east of the Sierran - Cascade axi s, from approx imately Mono County north to the Or egon border, altho ugh some popul ations are kn own from the Si erran crest pr oper, s uch as at D onner P ass or abo ve th e w est s ide o f L ak e Tahoe. It is also present in the higher elevat ions of the White M ountains. ELEVAT ION RANGE: From 5, 000 to 10, 000' elevat ion. Most po pulati ons occ ur betwe en 7,0 00 and 9,000' . HABITA T AND PLANT The species is typically found in Great Basin Sagebrush Scru b habitats COMMUN ITIES: on the east slope of the Sierra Nevada. In the White Mountains, it is predom inatel y as sociat ed wit h Mount ain Sa gebr ush Sc rub. PHENOL OGY : This species is str ictly univoltine. The adults fly from late Jun e through mid - August, w ith the majority of records in mid to late July. It o verwinters a s a first - instar larva. FOODP LANTS: N ative grasses used by this species remain to be determined. In the lab oratory, larvae can be readily r eared on Poa prate nsis L. b) C. o. near pallescens Emmel & Emmel DIAGNOS IS : Cercyonis o etus pallescens was descri bed by E mmel & Emmel (1971) from the Reese River Valley area of Lander County, Nevada, and is c haracterized by heavy white scaling across the ventral surface. In fresh specimens , the dorsal surface is also noticea bly li ghter than the col orati on in al l othe r su bspeci es, pre suma bly Unfinished book THE BUTTERFLIES OF CALIFORNIA by John F. Emmel, Thomas C. Emmel, and Sterling O. Mattoon [Document text truncated for crawler view.]