The floristic connection between the mediterranean region of Baja California and the Pacific islands of Alta and Baja California provides insight into the history and origin of the California Floristic Province. We present updated species lists for all California Floristic Province islands and demonstrate the disjunct distributions of 26 taxa between the Baja California and the California Channel Islands. These 26 plant taxa are found among the 16 Pacific islands without occurring on the intervening mainland of Alta California. Separate species lists for each island group (8 California Channel Islands and 8 Baja California Islands) were compiled. These lists were compared to the mainland California flora to identify species that occur on the California Islands and either the Baja California Pacific Islands or the mediterranean region of the Baja California Peninsula, but not the mainland of Alta California. This first compilation of the flora of the Baja California Islands and nomenclatural updates for the Channel Islands provide a platform for future research and conservation planning.
The islands of the California Floristic Province (CFP) are an important component of one of the world's great biodiversity hotspots (Myers et al. 2000). The CFP has commonly been referred to as an environmental island due to its regionally unique mediterranean-type climate, isolation (via oceanic and orographic boundaries), and corresponding globally significant levels of endemism. The plant community has had ample time to evolve since the emergence of wet winters and dry summers at least 5–10 Mya (Axelrod 1973) and perhaps earlier (Ackerly 2009, Keeley et al. 2011). The remarkable diversity and endemism of the CFP has been attributed to relative climatic stability during glacial-interglacial transitions of the Pleistocene (Lancaster and Kay 2012, Sniderman et al. 2013). The antiquity and distinctiveness of the California flora are well-recognized attributes that are accentuated on the islands of the Californias.
The degree of floristic similarity of 16 offshore islands, and their apparent role as refugia for species once present on the mainland, has been a persistent biogeographical puzzle. The CFP includes 16 islands from the adjacent coast of Alta and Baja California and extends south to the 28th parallel, including the CFP vegetation found on Cedros and Guadalupe Islands (Raven and Axelrod 1978, Moran 1996, Oberbauer 2002a). The CFP has an increasing gradient of aridity from north to south, but vegetation distributions are heavily influenced by fog presence; this fog provides additional moisture and reduces radiant loadings, significantly changing conditions for plants (Dawson 1998, Fischer et al. 2008, Vanderplank 2013). Physical data suggest that ocean currents have been stable for millions of years (Jacobs et al. 2004), which indicates the presence of fog during the development of the CFP.
Within the large number of plant species on the CFP islands and the mainland of Alta and Baja California is the occurrence of disjunct Baja California taxa on the California Channel Islands. These species are seen today only on the CFP islands and a distinct portion of the California mainland termed the Diegan Scrub, the San Diego region southward into coastal northwestern Baja California (Fig. 1; Axelrod 1978). However, the number and identity of this group of disjunct species has remained unknown.
To identify the species that define this pattern, we present (1) the first compiled checklist of the 8 CFP Baja California Islands, (2) nomenclatural updates to the flora of the 8 California Channel Islands, and (3) the plants that occur on the California Channel Islands and in the Diegan Scrub but not the rest of mainland California, hereafter referred to as “exemplar taxa.”
Development of Checklists
We assembled all existing species checklists for the 2 island groups (Alta California and Baja California) into 2 lists, then compared those lists with the flora of the California Floristic Province for both mainland Alta and Baja California to generate the list of exemplar taxa. The nomenclatural updates presented here are based on our best understanding of the island floras at the date of publication and follow the models of the sources indicated below for the 2 lists. Species now thought to be extirpated from the islands are denoted by an ampersand (&). Unvouchered taxa (including reports, photo vouchers, and personal communications) are indicated by a question mark (?). Nonnative species are indicated by an asterisk (*). When a subspecies was not recorded for all the islands and some ambiguity remains, the taxon is recorded at the species level only. Exotic species that were planted or are not naturalized (e.g., Nerium oleander) were deleted from checklists.
Baja California Islands Checklist
The most recent species lists for each of the 8 Baja California Islands (Appendix 1; Coronados, Todos Santos, San Martín, Jeronimo, Guadalupe, San Benito, Cedros, and Natividad) were compiled from the following data sources
The list of Jeronimo island flora, consisting of 8 taxa, was provided via personal communication from Steve Junak. We verified mainland Baja California flora distributions with the unpublished data of Jon Rebman and Bart O'Brien (personal communication 2013). Several taxa were added to the checklists based on recent collections, herbarium specimens recently encountered, and recent reports by Sula Vanderplank, Jon Rebman, and Steve Junak; these taxa are indicated by footnotes. The taxonomy for this checklist follows the Checklist of Baja California Plants in preparation by Jon Rebman.
California Channel Islands Checklist
The updated flora of the 8 California Channel Islands (Appendix 2; San Clemente, San Nicolas, Santa Catalina, Santa Barbara, Santa Rosa, Santa Cruz, Anacapa, and San Miguel) is based on the master checklist complied by Gary Wallace (1985). We included published updates to individual island floras found in the National Park Service checklist from Junak et al. (1997) for San Miguel, Santa Rosa, Santa Cruz, Anacapa, and Santa Barbara islands. For San Clemente Island we included updates from Ross et al. (1996) and for San Nicholas Island updates from Junak (2008). Additional taxa and island records are included based on recent collections on Santa Catalina Island by Sarah Ratay, observations and collections on San Clemente Island by Emily Howe (Soil Ecology and Restoration Group, SDSU), and herbarium specimens found in the Consortium of California Herbaria (CCH 2014), which are listed in footnotes. The nomenclatural revision of this checklist is consistent with taxonomy of the new Jepson Manual (Baldwin et al. 2012) through the use of the dynamic concordance tool provided on the Jepson e-flora website (Jepson Flora Project 2013). This checklist was developed to identify distribution patterns in the flora, and though the nomenclature has been updated, we have not confirmed identifications of questionable herbarium specimens, nor have we diligently pursued the lowest taxonomic rank beyond the published sources. We are also aware of additional updates to the flora of many of the Channel Islands in process by Steve Junak, including publication of the flora of Catalina Island; Junak's updates are not included here.
The identification of exemplar taxa was generated through a comparison of the 2 newly generated island checklists (Appendixes 1, 2) with the Jepson Manual (Baldwin et al. 2012) and the inventory of rare and endemic plants of CFP Baja California (O'Brien et al. 2014). A large number of species from CFP Baja California extend just slightly into southern California and are considered near-endemic to Baja California (O'Brien et al. 2014). As such, we include the near-endemic species of CFP Baja California as ‘absent’ in the mainland of Alta California (with the exception of Crossosoma californicum, which occurs on the Palos Verde Peninsula, and Euphorbia misera, which occurs in the South Coast region of California and beyond; Baldwin et al. 2012). These few species are consistent with the concept of Diegan Scrub as proposed by Axelrod (1978). Diegan Scrub is named for its occurrence in southern San Diego County and south into Baja California. It is a coastal scrub, rich in succulents, that includes many endemic and near-endemic species from CFP Baja; it also includes the coastal succulent scrub and coastal sage scrub ecoregions (González-Abraham et al. 2010).
The 26 exemplar taxa as they occur on 8 Baja California Pacific Islands and the 8 Channel California Islands.a CRPR column provides the California Rare Plant List ranking, and the BC column provides the State of Baja California Islands are listed left to right by increasing latitude. The Mainland column indicates continental occurrences. The protected rankings.
The California Pacific Islands in total contain 1239 unique taxa. The California Islands separately contain 976 taxa and the Baja California Islands 535 taxa. Twenty-six of these taxa have a disjunct occurrence between the Baja California Pacific Islands and the California Channel Islands, absent from Alta California outside the Diegan Scrub (Table 1). These exemplar taxa represent 16 families and 26 unique genera. No one growth form predominates; of the 26 taxa, there are 12 shrubs, 10 herbaceous annuals, 2 trees, 1 vine, and 1 xerophytic succulent (Table 1). Fabaceae is the largest family with 3 species on the exemplar taxa list.
These 26 taxa fall into 6 biogeographic subpatterns:
(1) multiple California Channel Islands and Guadalupe Island, 13 taxa
(2) Guadalupe Island-San Clemente Island only, 4 taxa (Hazardia cana, Leptosiphon pygmaeus ssp. pygmaeus, Lupinus guadalupensis, Phacelia floribunda)
(3) Diegan Scrub and the Channel Islands only, 2 taxa (Prunus ilicifolia ssp. lyonii and Salvia brandegeei)
(4) widespread, 1 taxon (Eschscholzia ramosa) occurs on 14 of 16 islands
(5) certain Channel and Baja California Islands and Diegan Scrub, 4 taxa (Bergerocactus emoryi, Crossosoma californicum, Euphorbia misera, Ribes viburnifolium)
(6) certain Channel and Baja California Islands, 2 taxa (Calystegia macrostegia ssp. macrostegia and Senecio lyonii)
The complex biogeography of the California Floristic Province can be explained in part by analyzing unusual floristic patterns seen on the California Pacific Islands. The 26 taxa shared by the Baja California and Channel Islands indicate a clear connection between these isolated and disjunct regions. What precludes these taxa from occurring on the Alta California mainland, and what does this suggest for the past extent of the CFP? Did these taxa evolve lo cally and then migrate, or does this pattern instead reflect a distant legacy?
Three current hypotheses attempt to explain the presence of these disjunct populations on the California Pacific Islands: (1) these species have the ability to disperse long distances yet for some reason have failed to establish on the California mainland; (2) these species were his torically more widespread, connected by former geologic linkages, and have been unable to persist on the Alta California mainland, remaining in refugia on the islands and in Baja California; or (3) the Channel Islands originally occurred farther south near San Diego (Atwater 1998) and as they moved north, fog-moderated climates retained species native to that region.
A number of the exemplar taxa have distributions on the Channel Islands and farther south in Baja California, suggestive of the refugium hypothesis (i.e., these species were once more widespread). Epling and Lewis (1942) proposed that the diverse Diegan Scrub has many taxa derived from Miocene vegetation that came from the north Mexican plateau. Axelrod (1978) states that Diegan Scrub was more restricted in the Pleistocene and expanded in the Holocene as arid-adapted taxa moved northward into the southern Channel Islands. Many CFP species are found in disjunct occurrences, scattered across Arizona and Mexico, not unlike the presence of these 26 species on the islands (Valiente-Banuet et al. 1998, Bhaskar et al. 2007). Small-scale topo graphic niches and facilitation by other species often support these occurrences. Axelrod (1978) also points out that the most important conditions for Diegan Scrub vegetation appear to be the absence of frost and the presence of a degree of summer moisture (in the form of fog). The cold California current and dense fog banks suggest that similarities in climate may help to explain the similarities in the floras (Fischer et al. 2008).
On the other hand, a number of the exemplar taxa are perhaps better explained as having evolved on the islands. This is especially true of many island Lotus species (Acmispon; McGlaughlin et al. 2011, McGlaughlin personal communication), Eschscholzia ramosa (Still and Potter 2013), and Crossosoma californicum (Wallace and Helenurm 2009), among others (Baldwin, 2007). For this pattern, we hypothesize local evolution and subsequent dispersal to similar niches on the CFP Baja Islands.
Guadalupe Island poses a particularly interesting study system. As the most isolated island, located roughly 270 km offshore, Guadalupe is a true oceanic island never connected to the Baja California Peninsula. Understanding the colonization of Guadalupe may contribute to understanding the dispersal patterns that we see today. Of 26 exemplar taxa, 21 of them occur on Guadalupe. Floristic similarities to the Channel Islands flora (at least 400 km north) may be due to the California current moving species from the California islands southward. Guadalupe Island has been above water for millions of years, and its current flora could represent the accumulation of species from the possible sources over this time.
In addition to a better understanding of the origin and evolution of the CFP, these species lists allow for improved conservation planning. Twenty-one of the 26 species are listed by the California Rare Plant Ranking (CNPS 2013) yet currently have no protection with the Mexican government. Twenty-five of the 26 species are listed in the rare plant ranking for Baja California (O'Brien et al. in press). These rankings are utilized in conservation planning in California and would be useful for land planning in Baja California. These combined islands species lists provide important information about the level of invasive species in the re gions. Of the 535 taxa on the Baja California Islands, roughly 14% (74) are introduced species (excluding planted species, which were not included in our analyses). In comparison, of the 976 species on the California Channel Islands, roughly 28% (278) are introduced species. Presumably, the lower proportion of invasive species on the Baja California Islands is due to less human use and fewer impacts to date.
The exemplar taxa identified by this process can now serve as the focal point of research to better understand the region's historical biogeography. Such research can incorporate phylogenetic relationships and their importance in community assembly (Webb et al. 2008). Functional trait analyses would help determine if these taxonomically diverse species share inherent traits that explain their disjunct distributions. Comparative phylogeographic studies of these shared taxa would test the specific hypothesis discussed above, among others, and offer a glimpse into the past. Additionally, the knowledge of patterns and modes of dispersal would provide valuable natural history information. Further genetic work to resolve the origin of these island specialist species would help to elucidate questions of timing, and definitively separate neo- and paleoendemics in the exemplar list.
Conservation activity in Baja California has recently expanded, yet knowledge of plant distributions and ecology remains limited. We hope that these updated checklists of the California Pacific Islands will facilitate species status and distribution updates on both sides of the border, and foster well-informed conservation decisions that promote expanded science and management for the transborder ecoregion.
This work is based on the legacy of botanists who have explored and documented the floristic diversity of California and the Pacific Islands. In particular, we thank the following individuals who have directly aided this paper. Steve Junak's expertise has been invaluable; we are most grateful for his personal communications and guidance. His dedication and role as an information clearinghouse for the island floras is vital for islandfocused plant work. We are extremely grateful to Jon Rebman of the San Diego Natural History Museum (SD) for his assistance with the Baja California Island checklist, as well as his additional data, nomenclatural reference works, and taxonomic assistance. Karen Rich and Judy Gibson at SD also provided information. Bart O'Brien and coauthors kindly gave us access to a great deal of informative data in their in-press manuscript. Two anonymous reviewers contributed greatly to the quality of this manuscript.
Plant species checklist for the 8 Baja California islands (Cedros [CED], Coronados [COR], Jeronimo [JER], Guadalupe [GUA], Natividad [NAT]; San Benito [BEN], San Martín [MAR], and Todos Santos [TOS]). An ampersand (&) indicates an extirpated species; a question mark (?) indicates an unvouchered taxon; and an asterisk (*) indicates a nonnative species.
Plant species checklist for the 8 California Channel Islands (Anacapa [ANA], San Clemente [CLE], San Miguel [MIG], San Nicolas [NIC], Santa Barbara [BAR], Santa Catalina [CAT], Santa Cruz [CRU], and Santa Rosa [ROS]). An ampersand (&) indicates an extirpated species; a question mark (?) indicates an unvouchered taxa; and an asterisk (*) indicates a nonnative species.