The case for using geography to understand phenology

 Alexandra Permar and Conor Flynn collaborated on this project.

Calochortus is an astonishingly diverse genus of flowering monocots, with common names that include Mariposa lily, Sego lily, and globe lily, as well as pussy ears, fairy-lantern, star-tulip, and others.  Their center of diversity is in CA, which has 48 out of the 78 total species in the genus.  In Arizona, there are 6 species, ranging in color from white, to pink, to yellow, and sometimes orange, with diagnostic darker markings on the inner base of the petals.  

    

    This genus in Arizona makes an interesting case study of biogeography and phenology because each species occurs in a relatively distinct (but overlapping) elevation and geographic location.  Also, because the flowers are very showy, there are more than 2,400 Research-grade observations documented on iNaturalist.   

    The plant grows quickly from underground tubers in the spring, flowers quickly, and then dies back; the growing stem and then the seed heads are relatively inconspicuous and are rarely photographed.  This means that almost all of the iNaturalist observations are of the flowers, which makes studying the phenology easier because there is less work needed to annotate images as flowering or not flowering.  Nevertheless, we still reviewed observations and annotated and removed any photographs that didn't show flowers from the phenology analysis below.  

Methods 

Research-grade Calochortus observations in AZ were exported from iNaturalist and processed in Excel and graphed in Excel and Tableau.  iNaturalist Research-grade observations are used for research, but observations with inaccurate locations can lead to misleading analyses and must be removed.  

A total of 2350 observations were exported.  Taxa were classified by species (even though subspecies IDs exist for several hundred observations).  Cleaned records with geopositional accuracy > 1000 m (excluding blanks; there are 594 blanks).  Removed 61 records with geoprivacy obscured. (In iNaturalist, obscured geoprivacy impacts the geopositional accuracy of an observation to +/- 500 square kilometers.)  8 observations were noted as not flowering and were removed from analysis.  After cleaning, the database contained 2066 records (observations).  This shows that about 88% had location precision that met our inclusion criteria.

 Calochortus kennedyi had the most observations (782),  C. ambiguus and C. flexuosus tied for 2nd most observations (534), while C. aureus (148) and C. nuttallii (61) had the least number of observations. 

Geocoding Elevation

iNaturalist does not record elevation of observations, so it is necessary to intersect the observation points with a Digital Elevation Model in GIS.  Observations were added to ArcGIS Pro and displayed using their XY coordinates.  Connect to elevation in Esri AGOL Living Atlas : Ground Surface Elevation - 30m (image service / raster DEM).  Use Geoprocessing tool Extract Multi Values to Points ( Need Spatial Analyst license).  Copy resulting data back to excel. 

        Tableau Public was used to graph data.  Calochortus Bio-Geo-Phenology | Tableau Public

Results: Biogeography and Elevation

    Ordered by average elevation, Calochortus kennedyi and C. flexuosus are the lowest elevation species, C. aureus, C. ambiguus, and C. nuttalli occur at middle elevations, and C. gunnisonnii occurs at the highest elevations.  

iNat allows visualization of biogeographic comparisons.  (Or new compare tool) 

C. kennedy (blue) is clearly visible at lower elevations along the foothills of the AZ mountains, and then extends to the West beyond the distribution of any other species.  C. flexuosus (green) is also visible to the NW of AZ, extending through central AZ in the lower elevation valleys of the Verde, Salt, and Gila rivers.  C. gunnisonnii (purple) is clearly visible in Colorado, barely extending down to the high country in AZ.  C. aureus (pink) stakes out a unique territory on the Navajo nation in NE AZ and SE Utah. C. ambiguus (red) is found throughout the mountains and high country of AZ.  

 Graphing both latitude and elevation help to illustrate this biogeographic comparison.  In the graph below, it can be seen that C. kennedyi is the most southerly of our Calochortus, and occurs at the lowest elevations.  C. flexuosus occurs farther north, but also at low elevations.  C. ambiguus marks out a consistent territory at higher elevations than other species, depending on elevation. Gunnisonii is a clear outlier, occuring only at high elevation in our study area.  There is a fair amount of overlap between C. aureus and C. nuttallii.

        This biogeographic comparison can help differentiate similar-looking species.  There are 4 white Calochortus in AZ.  C. gunnisonni is clearly only a high elevation species, although C. ambiguous can also occur at high elevations.  In the northern part of the state, C. flexulosus occurs at lower elevations, C. nuttallii at middle elevations, and C. ambiguous only occurs at the highest elevations. 

Results: Phenology

Overall, for Calochortus species in AZ, flowering begins at low elevations around week 13 (last week of March) and progresses to higher elevations, generally wrapping up around week 29 (third week of July).  Flowering species composition changes from C. flexuosus and C. kennedyi to C. ambiguus, with C. aureus and C. nuttalli thrown into the mix.  C. gunnisonni (barely visible at the far upper right) is, of course, the last to flower.  This chart also highlights how C. aureus is mainly restricted to 1600-1800 meters ( 5200-5900 feet).  

    Calochortus ambiguus shows the strongest relationship between elevation, latitude, and flowering date.  Based on this scatterplot, we were able to confirm that the location is not accurate for the observation at the far left side of the plot. 

Calochortus kennedyi, C. aureus, and C. flexulosus do not show as strong of a pattern, and C. gunnisonnii did not have enough observations in AZ to analyze.  

Example of C. kennedyi:

Graphing flowering week of C. kennedyi separately for Latitude and for elevation shows that there is some pattern, but it is clearly quite week.  The R squared values for these charts are 0.22 and 0.12.  Interesting, it looks like flowering actually starts at middle elevations (around week 11), then flowers appear at lower and higher elevations (week 16), until finally only the higher elevations are still flowering (week 20).  C. aureus and C. flexulosus show somewhat similar patterns (data not shown).  

Discussion

    There are many possible ecological and data collection reasons for weak r values:

1) Micro-site variation: south-facing aspect may have more impact than elevation.  Whether a site is forested, if there are shadows from cliffs, etc.

2) Genetic variation:  if a single species showed up with 2 distinct populations, that would be evidence for possible speciation or subspecies, but all populations showed continuous variation.  There may still be genetic variation between meta populations, or within single individuals.

3) Flowering period not known: our records only show flowering presence, not the start of flowering or the total period of flowering.  Correlations might be better if we had complete data, but our data is not complete: we only have point observations.  For example, if species A flowers from April 1-30 and Species B flowers from April 28-May 28, a visitor on April 28 would record both species as flowering at the same time, even though they do have distinct flowering periods.  

Joshua Trees Flower Episodically in Arizona

Intro

Joshua trees are large, visually-striking trees in the Mojave desert.  Most of them are in California and Nevada, but there is also a population in the western part of Arizona.  They are frequently photographed and there are more than 17,000 observations (1,000 in AZ) saved on iNaturalist, a website and app used to document biodiversity.  

Arizona Joshua Tree locations observed on iNaturalistiNaturalist.

Joshua trees produce showy clusters of white flowers at the tips of their branches in March, but this year I noticed they were not flowering.  I wondered how often they produce flowers and decided to answer this question using data from iNat.

53 observations in Arizona showed evidence of flowering, always in March and April.  

Methods

iNat observations of Joshua Trees (Yucca brevifolia) in Arizona were marked using the Plant Phenology option in the Annotation Field.  Plant phenology (flower budding, flowering, or fruiting) was determined based on visual inspection of the saved photos in iNat.

The iNat website has a nice phenology visualization tool.  This view is filtered to show only Arizona observations.  However, it cannot show differences in phenology from one year to the next.  

Observations were then filtered on the iNat Explore page by adding &term_id=12&term_value_id=13 to the URL and downloaded for analysis in Excel.

More info about using iNat search URLs.

Results

I graphed the results starting in 2017, because there are fewer iNat observations before 2017.  It appears Joshua Trees flower episodically.  According to this data, Joshua Trees flowered in four out of the last 7 years.  They appear to alternate years from 2017-2020, but then skipped 2021 and flowered in both 2022 and 2023.  

It is interesting to note that the Arizona Joshua Trees didn't flower in 2020, which was one of the wettest springs in the last 10 years. I wonder whether the spring moisture determines flowering, or if perhaps other climate variables, such as moisture in the fall, are more important.  Another possibility could be that the trees are only able to flower every other year, and that the trees were inhibited from flowering in 2020 because of the large number that flowered in 2019.  

According to this data they were able to flower in both 2022 and 2023, but at reduced numbers both years.  Perhaps the plants flowering in 2023 were different from the trees that had already flowered in 2022?  I can't answer that question with this iNat data.  While it appears that the areas of flowering in 2022 and 2023 were the same, not all of the Joshua trees in a given area necessarily flower, even in good years...

Where is most biodiverse?

 iNaturalist can be used to compare biodiversity in different locations.  

To do this, it is important to allow for different search effort in different areas: for example, a search of the middle of the Amazon might not return many species, not because there aren't many, but because people haven't observed them.

For the U.S., we will assume that search effort is at least somewhat comparable.  We will also look at total observations and divide them by total species to get some idea of the effort required to document a new species in that area.

We decided to investigate my home town in Washington State, Alexandra's home town in Ohio, one of our favorite places: Maui, and our current home in Arizona.

Port Orchard, WA

A 100 km search radius includes most of Puget Sound, the Olympic mountains, and Seattle.

Dublin, OH

A 100 km search radius includes most of central Ohio creeks, woodlands, agricultural lands, and all of Columbus OH.

Prescott, AZ

A 100 km search radius includes canyons and mountains between Flagstaff and Phoenix.

Maui, HI

A 100 km search radius includes all of Maui as well as ocean channels and nearby islands.

Summary Table

According to this analysis, Washington is most biodiverse, followed by Ohio, then Arizona, and in last place is the island paradise of Maui.  

Island biogeography teaches that islands, although unique for their level of endemic plants and animals, should be deficient in overall biodiversity if there hasn't been enough time for biodiverse taxa to colonize or evolve on the island.  This seems to be the case with Maui, which is especially depauperate in insects, given its tropical location.

Arizona, despite its diverse environments, is significantly lacking in Fungal diversity, which could be due to the arid climate.

The American Midwest, with its perfect seasonal growing climate, is remarkably biodiverse, and would probably surpass Washington if this search had included a more varied part of the Midwest with more mountains and/or rivers and lakes.

Washington tops the list as most biodiverse, probably because of the presence of diverse environments, with everything from alpine mountain tops to ocean kelp forests, and everything in between.  This is despite the lower number of insect species compared to a lower latitude place like Ohio.

A caveat to this entire analysis is revealed by the Total Observations in each area and the resulting Observations per Species.  While I had assumed that search effort would be equivalent across the US, it is evident that many more people are using iNaturalist around major cities like Seattle and Columbus.  Arizona is more sparsely populated and has many less observations, and Maui, despite the huge numbers of tourists, is even less populated.  This means that is only takes 20 or 27 new observations (on average) to observe a new species in Maui and Arizona, respectively, while it takes 47 or 72 new observations in Ohio and Washington, respectively, to observe a new species.  So even though Washington has the most observed biodiversity in iNaturalist, it may be easier to find new, previously-unobserved biodiversity in Maui or Arizona.

Seed Zones for Western Wheatgrass

The USDA Pacific Northwest Field Station has published research examining the population geographic structure of Western Wheatgrass.  They measured three major phenotypic factors from grass seed collected throughout the interior northwest: height, flowering date, and leaf width.

Sunflowers in NM

Common sunflowers in NM:
Perennials:

Helianthus ciliaris Blue-green, short, forming extensive colonies from perennial roots.  Leaves sessile.

Annual (no rhizomes):

Helianthus annuus  -Phyllaries ovate, abruptly narrowed to acuminate tip, ciliate

Helianthus paradoxus  phyllaries lanceolate, gradually tapering to the tip, not ciliate.  Usually no branches, no white “eye” in flower center. In wetlands.

Helianthus neglectus  phyllaries lanceolate, gradually tapering to the tip, not ciliate.  Usually branched, with a white “eye” in flower center.  Lower leaves w/ cordate bases.  Larger heads than H. petiolaris. 

Helianthus petiolaris  phyllaries lanceolate, gradually tapering to the tip, not ciliate.  Usually branched, with a white “eye” in flower center. Lower leaves w/o cordate bases.

Key based on Allred's Flora Neomexicana

Thistles in NM

I've compiled a list of common Cirsium species in the lowlands (not mountains) of Southern New Mexico, and mapped their collectioned vouchers from across NM.

Map created using swbiodiversity.org 

Cirsium ochrocentrum – common.  Flower heads are bigger, and petals are longer than C. neomexicanum, often more brilliantly-colored, but plant is smaller.  Flowers usually taller than wide. 

Cirsium neomexicanum  -- Flower head looks more “bushy” because spines are more strongly spreading than on C. ochrocentrum, allowing flower to open more (wider than tall)

Cirsium texanum  - like C. ochrocentrum, but flower heads less than 25mm long, smaller overall.

-Based on Allred's Flora Neomexicana

Passion-flower Distribution in the Sonoran Ecoregion

Source: SEINet

Passiflora (Passion flower vine) is a tropical genera of vines, which reach their northernmost distribution with P. mexicana in the Santa Teresa mountains, North of the Gila River.  They produce edible passion fruits, but the herbiage is extremely unpalatable to animals, to the point that (reportedly) starving horses will not eat P. foetidius due to the stinky sticky hairs.

I have never seen these monsoon-bloomers in the wild in Arizona, but hope springs eternal in the Sky Islands...there are three species in Arizona:  P. arizonica, P. bryonoides, and P. mexicana:

P. arizonica ca 4-5.5 cm in diameter, whitish, the corona white or purplish.

P. bryonioides ca. 2.5-4.5 cm in diameter, whitish with purplish bands on corona. 

P. mexicana ca. 2-3 cm in diameter, light green or yellowish green, the corona red or reddish purple.

There are two more species in the Sonoran Desert south of the border:

P. palmeri (no description available)

P. suberosa (no description available)

A Range Overlap of the Pinyon Pines in central AZ

In which mountain range could one find three different U.S. Pinon Pine species (P. cembroides, P. edulis, and P. monophylla)?

Trick question!

Surprisingly, not a major mountain range, but the low hills surround Aravaipa Canyon.  This area may be an important biogeographical linkage between the sky island mountain ranges to the south, dominated by Mexican Pinyon (Pinus cembroides) and the larger mountains to the north, dominated by Pinus edulis.  Pinus monophylla is more of a Colorado plateau and California mountain species, but with weird disjunct populations as far South and East as the Florida mountains near Deming, NM!  

Specimen collection data from SEINet.

Four of my Favorite Characteristic New Mexico Shrubs

Chilopsis linearis, Desert Willow.  In landscaping, this large-flowered shrub will flower for almost half the year.  It is a dependable riparian tree that can survive in washes that are too hot and dry for cottonwood.

Forestiera pubescens.  New Mexico Privet.  Native shrub in the Olive family can form thickets at the base of canyon walls and along upper benches above rivers.  Birds prize the berries almost as much as the non-native Russian Olive.

Lycium pallidum.  Pale Wolfberry.  Lyciums are widespread in the Southwestern deserts, and Lycium pallidum is the most widespread.  Goji berries are Lycium barbarum (nonnative), and Lycium pallidum has been cultivated for its fruit for 1,000s of years.  The trumpet-shaped flowers attract Sphyngid Hawkmoths and Hummingbirds.  

Robinia neomexicana.  New Mexico Robinia, Fabaceae.  Despite its rapacious thorns, this plant made my list for its tenacity and abundance across a range of habitats.  From riparian area in the South to Ponderosa hillsides in the North, Robinia provides abundant pink blossoms and herbaceous cover. 

In what mountain range could one see four different species of Juniper trees?

 These maps of specimen collections are from http://swbiodiversity.org/seinet/