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Introduction

California is a varied state noted for its diverse This book was written with these ideas in topography, geology, plant communities, and mind. The families I have selected are those 50 native plants. Of the 173 or so native and natu- important ones plus several others that help de- ralized vascular plant families, there are in ex- fine our vegetation. For example, maples (family cess of 5,800 species, around 1,000 of them in- Aceraceae), California buckeye (family Hippocas- troduced from other parts of the world. That is tanaceae), California bay (family Lauraceae), and an impressive figure for an area our size. alders (family Betulaceae) are such key compo- How can a naturalist make sense of this nents of woodlands and forests that I felt com- tremendous diversity? The first step is to look at pelled to include them even though each family the big picture by learning to recognize plant has only a few species. families. Despite the intimidating number of Because there is already an excellent book those families, roughly 40 to 50 families con- on major desert families (California Desert Flow- tain more than 85 percent of our flora. Learning ers: An introduction to families, genera, and species to recognize these keystone families goes a long by Sia and Emil Morhardt, published by the ways toward the process of identification and al- University of California Press), I have tried to lows you to create a framework for most of the avoid delving into detail on desert plants, al- genera and species you are likely to encounter. though it has been impossible to exclude them Although there are many versions of keys to all. The major families selected for this book these and other families, learning the major characterize the area we refer to as the Califor- field characteristics (often with the aid of a good nia Floristic Province, which also coincides with hand lens) saves a lot of time and makes the a geographical realm known as cismontane Cal- process more enjoyable. Close observation of ifornia. Just what is meant by those terms? habit, leaves, flowers, and fruits leads to impor- Floristic provinces are assemblages of plants tant information that is useful in other ways. that belong to plant communities typical of re- For example, it helps to explain pollination, gions that have a similar overall climate and adaptations to habitats, and life cycles. geographical coherence. The plants in each

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province form repeatable associations that show more dramatically than at any previous time in adaptations to their home and differ, sometimes history. Despite the inconvenience of a classifi- dramatically, from plants from other floristic cation in a state of flux, think of these changes provinces. The California floristic province is one as an exciting challenge to learn more about the of the most distinctive in the world and repre- evolutionary relationships of all organisms. sents one of five major areas with a Mediter- Certainly these changes are frustrating to those ranean climate—cool to cold, wet winters and wishing to learn an unchanging system, but warm to hot, usually dry summers. This province that is not the reality of what is going on. extends from the Rogue River in southwestern The reason for this state of affairs is that Oregon south through California into the north- many important lines of research are creating a western fringe of Baja California. Most of it lies more complete picture of how species and gen- to the west of the main mountain crests of the era are related, and which families they belong Klamath Mountains, Sierra Nevada, Transverse to. In addition to the more classical lines of in- Ranges, and Peninsular Ranges, a high backbone quiry such as external form (morphology), that delineates much of California. This region is anatomy, chromosome studies, and details of also referred to as cismontane, that is, the region pollen, the growing arsenal of information in- on the ocean side of the mountain crests. The cludes studies of biochemistry involving pig- transmontane region to the east is mainly desert ments, poisons, perfumes, proteins, and many and supports other floristic provinces. other compounds. Add to this the rapidly ex- As with other aspects of nature, the lines be- panding field of DNA studies that plot the rate of tween provinces is often blurred so that various changes in selected genes, and you have a far desert elements and families enter the coastal richer and more detailed story of true relation- mountains of Southern California. As a conse- ships. Computer-generated cladograms that dis- quence, I have included certain families such as play the degree of relatedness of plant groups the cactus family (Cactaceae) and spurge family present an evolutionary scheme displayed as a (Euphorbiaceae), which are best represented in branched system. These cladograms are used to our deserts. In this endeavor, I have tried my determine the limits of families and genera. best to be even handed and to balance impor- Consequently, several well-known and tant plants from all parts of the state. widely studied families are now in a state of change. I wish I could say that these changes IMPORTANT PLANT FAMILIES IN will be permanent and all you need do is learn CALIFORNIA the new classifications, but as we continue to learn more and examine a larger array of genes, California has around 173 different vascular we will not only refine the now-current system plant families, some native and others intro- but also make additional changes. duced. The introduced ones may appear to grow Some long-recognized families have been on their own in natural habitats. Many of these split into two or more separate families while nonnatives are accidental introductions brought others have been “lumped” together. Still others in on bricks or ballast, as contaminants in culti- are so complex that totally new alignments are vated crops, on domestic livestock, and by hu- being made. mans. Other plants have “escaped” from gardens My approach in this book is somewhat con- and cultivated fields to grow on their own. Many servative; because the current Jepson Manual is of these are invasive and seriously threaten the still the standard reference for workers and stu- diversity of our native flora. dents and because so many other local floras We live in a time of rapid change in the follow a similar system for the families, I have world of biology. It is important to know that continued to use many of the familiar family the state of defining plant families is changing definitions. But I also alert you, the reader, to

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changes that are proposed or have been ac- Douglas-fir (Pseudotsuga menziesii) for a tree cepted; future field books will eventually reflect that despite its very different cones and bark re- these changes. minded someone of a fir (Abies spp.). I have deviated from the older system(s) Not every plant or plant family has a well- with the large, important lily family (Liliaceae), established common name, but since people want which is now considered to consist of many a common name, I have tried to provide them separate, sometimes unrelated families. Al- when possible. Not everyone will agree with my though I have not treated all of these splinter choice of common names, and some will dis- families here, I am describing some of the agree with my use of one common name over major ones that are easily recognized: the agave another. Where there is more than one well- family (Agavaceae), onion family (Alliaceae), known common name, I have included it. and brodiaea family (Themidaceae). Scientific names are based on carefully crafted rules for naming, and are in a latinized HOW PLANT FAMILIES, GENERA, AND form. Because scientific names can be recog- SPECIES ARE NAMED nized by scientists throughout the world re- gardless of the language they speak, the names All plants have names, and it is the name we give a real sense of permanence and are imme- turn to first when we want to learn more about diately recognizable anywhere. a particular group. You will find that plants bear two kinds of names: trivial or common names PLANT FAMILIES that are in everyday use by the average person, and scientific or Latin names, that are used Because this book focuses first and foremost on throughout the horticultural and botanical families, I will talk first about the rules for scien- world. There are advantages and disadvantages tific family names. (Families are larger than to both kinds of names, but for greater preci- genus and species, often embracing several dif- sion, scientific names are preferred. Common ferent genera and many species, although some names are not always standardized, and many families are very small.) All family names end in plants and plant families—for example, Fa- -aceae and are based on a type genus: for example, gaceae, aka the oak or beech family—have more we have Rosaceae (rose family) based on the type than one common name. Common names may genus Rosa; Liliaceae (lily family) named for the also allude to relationships that do not exist. For type genus Lilium; and Orchidaceae (orchid fam- example, the evening-primrose family (Ona- ily) from the type genus Orchis. Those examples graceae) and primrose family (Primulaceae) are are based on names that are cognates of common not at all closely related; evening-primroses and English names, but many family names are not their relatives belong to a different evolutionary recognizable, such as Ranunculaceae (buttercup line. Perhaps the first people to notice this family), Rhamnaceae (buckthorn family), and beautiful family were struck by the showy flow- Scrophulariaceae (figwort family). Understand- ers that they imagined looked like oversized ing the derivation of these names makes learning primroses, but botanically, the two families dif- them much more enjoyable. Ranunculaceae fer by many traits (turn to pp. 000 for a de- refers to little frogs, because many buttercups live scription). When these sorts of common names in wet areas, the habitats for frogs. Scrophulari- have become so embedded in the language that aceae is named for the type genus Scrophularia, they are permanent, the name is hyphenated to based on the belief that it cured the skin disease indicate that it represents a special combina- known as scrofula. tion. So we have corn-lily (Veratrum spp.) for a It is also important to understand that sci- perennial wildflower that belongs to the lily entific names may change as more is learned family but is not a true lily (Lilium spp.), and about a particular group. For example, there is

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considerable controversy over what belongs to Both genus name and specific epithet are un- the lily family (Liliaceae). Research indicates derlined or italicized. Although the initial reac- that there are several separate evolutionary tion of the novice may be that scientific names lines in this family if you define it to include the are impossibly difficult, in fact English speakers broadest possible concept. Each line is often have a decided advantage since many of the lat- given a separate family name. But if you look at inized names have cognates in English. A few the bigger picture and choose to focus on the examples of recognizable scientific names fol- common ancestry to these evolutionary lines, low: Lilium maritimum (coast lily)—the genus you may prefer to lump all the species together name is the Latin version of lily, followed by a into one very large, inclusive family. word that means coastal (maritime). Viola pur- I have usually followed the family concepts purea (pine or oak violet). The name Viola al- given in the Jepson Manual but as commented ludes to the violet color of many flowers in the on above, new DNA research is turning many genus, although this particular species happens families topsy-turvy. In short, evidence from to be yellow; purpurea means purple, perhaps this research is causing an almost unprece- because the underside of the mature leaves is dented reorganization of families. The Liliaceae purple. Delphinium nudicaule (scarlet larkspur): mentioned above is now sliced into more than Delphinium comes from the Latin word for dol- 16 separate families (not all of these are Cali- phin and is a cognate of our English word— fornian). This reorganization is highly relevant larkspurs have a sleek, streamlined outline that to those studying evolutionary relationships but is reminiscent of the body shape of dolphins; it raises havoc with conventional books for the nudi means naked (nude), caule stem (think of amateur botanist and nature lover. the word cauliflower, meaning stem flower).

PLANT GENERA HOW TO USE THIS BOOK Besides families, this book covers many impor- tant genera and even some species. After family If you already know the name of the plant but comes genus (plural genera). A genus consists are curious about the family it belongs to or are of closely related kinds of plants. For example, simply trying to organize your knowledge bet- among wildflowers we have violets (genus ter, turn directly to the main section of the Viola), daisies (genus Erigeron), Indian paint- book, where each family is listed alphabetically brushes (genus Castilleja), buttercups (genus by its scientific name. If you know only the Ranunculus). Some genera have only one common name of the family, consult the index. species; others have dozens or even hundreds. If you have a plant to identify and have no idea The kinds of violets or daisies are what we call what it is, refer to the family key (below). species (singular and plural are the same). Cal- CALIFORNIA PLANT FAMILY ACCOUNTS ifornia is blessed with many species of violets, including V. douglasii (Douglas’s violet), V. pe- Each family entry starts with a short statement dunculata (wild pansy), V. adunca (dog violet), V. about recognition at a glance—what to look for macloskeyi (white meadow violet), V. ocellata as a first step to identifying the family. When (western heartsease), and many more. this does not suffice—or if you are anxious to learn more—a series of descriptions expands on PLANT SPECIES other family traits including the habit, leaves, Each scientific species name consists of two branches, flower shape and arrangement, flower parts: a genus name, given first and capitalized; parts—numbers of sepals, petals, stamens, and and a specific epithet, given second and starting pistils—and fruits. The term habit describes the with a lowercase letter. (Think of how most form of a plant, such as shrub, tree, herbaceous people use two names to identify themselves.) perennial, annual, or bulb.

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Because many families look alike or are If you are new to keying, here are some closely related, there is a section dealing with pointers to bear in mind: similar-looking families (which are not always closely related despite a superficial resem- • Keys are imperfect because nature is not al- blance), with quick suggestions to distinguish ways consistent. them from the family you are learning. • Be sure to note as many features about the The statistics entry alerts you to the distribu- plant as possible if you do not have a fresh tion, habitat preferences, size, and economic specimen in hand. You will want to note the uses of the family. leaf shape, arrangement, and form (simple Finally, there is a section on California gen- or compound), the habit of the plant (herb, era and species. For small families, I have tried shrub, or tree), the arrangement of the flow- to be as inclusive as possible so that, for exam- ers, and the details of the flowers and their ple, you will learn about all four tree species parts, including shape and any special fea- that belong to the maple family (Aceraceae). For tures. If your specimen has fruits or seeds, large families with dozens to hundreds of those should also be noted. species, a comprehensive breakdown of the • Write down the choices you have made so family is beyond the scope of the book, but I that when a particular choice is not clear have included a fair sampling of genera and you can backtrack if the family you choose species that I consider typical, common, or in- does not fit the description of your plant. teresting. These examples are presented under • Try working the key backwards, starting logical categories that act as a guide or simpli- with a family whose name you know. By fied key to separate the given genera. following the steps that lead to the family CALIFORNIA PLANT FAMILY KEY name, you will learn more about how the key works. The key is designed to minimize technical fea- • Practice makes perfect, and the more repe- tures, although many details such as types of titions you do, the easier the process fruits and the ways anthers shed their pollen are becomes. necessarily mentioned. A good 10x hand lens is a valuable and necessary adjunct to see many of • Consult the glossary at the back of the book these traits. I often include more than one trait for terms that are not familiar. Learning for each step in the key but sometimes the sec- these terms is like learning a new language. ond trait given is not necessary to successfully • Consult the illustrations of flower parts lo- use the key. (Second or third traits at each step cated in the glossary. are there to help confirm your decision.) Although I have attempted to take into con- As you peruse family traits you may find cer- sideration exceptions to the general family char- tain unfamiliar terms. Technical terms are use- acteristics, there will doubtless be some that I ful when they replace an otherwise sentence- have missed; the key should work to identify a long description. The glossary will help make plant to family most of the time. sense of these terms. Be sure to study the line If you have a plant to identify, you will want drawings to see how the language translates vi- to use the simplified key and carefully follow sually. the choices at each step. To start keying, choose Similar-looking families may seem confus- one of the three groups—conifers, monocots ing to separate. (Examples include the borage and monocotlike plants, or dicots—then turn to and waterleaf families, the morning glory and that group and proceed from there. Most steps nightshade families, and the onion and brodiaea in the key have two choices but a few have three families.) In such cases, check the description of or rarely more. the look-alike family. I comment on families

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that might be confused with one another and packed together in a head that resembles a sin- compare them. gle flower; parsleys bear many tiny flowers in Finally, note that no single trait ever serves compound umbels. By contrast, the Rosaceae as a sure means of recognizing a family; rather, (rose family) varies in just about every feature you need to look at a combination of traits. For from leaf design to flower arrangement, from example, knowing that the Onagraceae plant habit to stamen number, and from habi- (evening-primrose family) is characterized by tat to ovary position. Yet, even this large family an inferior ovary does not allow you to separate offers important clues to its identity such as it from other families with inferior ovaries. But the presence of a hypanthium, a single-rose- when you note that your plant also has four like flower design, specific types of fruits, the petals, four sepals, four or eight stamens, and a frequent presence of stipules on the leaves, capsule-type fruit, you can be confident that the and sepal-like bracts that alternate with the plant belongs to this family and no other. true sepals on flowers of the nonwoody Families sometimes differ in their variabil- species. ity. For example, the Asteraceae (daisy family) Remember: Practice makes keying go faster and Apiaceae (parsley family) are intuitively and more smoothly. Use this book often and easy to recognize because of the appearance of you will grow more proficient at keying and their flowers: Daisies have many flowers find satisfaction in your new skills.

6 introduction