April 22, 2021

Stemless White Violets in Pennsylvania

Gosh they're confusing. Here's a summary table derived from the master key on Harvey Ballard's website. These are only notes; please cite his website/papers, not this journal entry, which is just a PA-centric re-formatting, including many direct quotations. Again, this is not my work.

Viola odorata can also have white petals sometimes, but it's usually pretty easy to distinguish from this group based on its style shape and large flowers.

Viola... lanceolata primulifolia renifolia minuscula blanda incognita
leaf blades
length vs. width
3 – 8× 1.5 – 2× < 1.2 × < 1.2 × < 1.2 × < 1.2 ×
base cuneate rounded – subcordate deeply cordate shallowly to deeply cordate deeply cordate shallowly, broadly cordate
carriage** lying on substrate
(or widely spreading)
elevated and spreading elevated and spreading elevated and spreading
color bicolorous
(darker green above)
faces uniformly green bicolorous
(darker green above)
bicolorous
(darker green above)
pubescence entirely glabrous
or
dense on one or both surfaces
strictly glabrous glabrous except for scattered appressed hairs on upper surface glabrous or sparsely to densely hirsute, including petiole
shape broadly ovate or suborbicular to broadly reniform, occasionally suborbicular narrowly to broadly ovate, deltate-ovate, suborbicular or subreniform (very rarely reniform)
apex abruptly apiculate, obtuse or rounded
at least some broader than long
apex obtuse to acute-acuminate
at least some broader than long
apex obtuse to acute-acuminate
margins usually shallowly crenate shallowly crenate low-serrate, teeth noticeable
sinus length < ¼ length of blade > ¼ length of blade
(often > ⅓)
basal lobes touching or overlapping well-separated or divergent
petioles red-tinged or spotted greenish
rhizome vertical, twisted
lacking stolons*
horizontal, stoloniform
producing stolons*
horizontal, stoloniform
producing stolons*
horizontal, stoloniform
producing stolons*
lateral petals beardless (or nearly so) beardless (or nearly so) beardless bearded
cleistogamous capsule
coloration
purple-spotted unspotted or with fine red spots red-purple to purple spots/blotches purple spots/blotches
peduncle short and prostrate
arcing upward only when opening
long and erect, often surpassing petioles prostrate, then arcing, sometimes surpassing petioles prostrate, then arcing, much shorter than petioles
length /mm 3.5 – 8 7 – 14
seeds
color
orange-brown – brown
unspotted
med- dark- olive-brown or brownish-black
fine black spots
brownish-blackish
unspotted
light to medium brown
unspotted
dimensions /mm 1.7 – 2.4 × 1.2 – 1.4 0.8 – 1.4 × 0.7 – 0.8 1.2 – 1.3 × 0.7 – 0.8 1.6 – 2.2 × 1.0 – 1.3
*stolons may be above or below surface of substrate, and may not be apparent until summer. **in life, especially during fruiting

Synonymy with the taxa currently on iNat:
V. minuscula V. blanda V. incognita
V. macloskeyi (in part)
V. pallens (in part?)
V. pallens ssp. pallens
V. pallens ssp. subreptans
V. macloskeyi ssp. pallens
V. blanda (in part)
V. blanda var. blanda
V. blanda (in part)
V. blanda var. palustriformis

Posted on April 22, 2021 00:52 by ddennism ddennism | 0 comments | Leave a comment

January 13, 2020

How many cultivated plants are marked as such on iNaturalist?

Thought I'd try to address this question with a semi-scientific approach:

For the past year (1/12/2020 - 1/12/2019) in a 25 square-mile square centered on Blacksburg, VA:

To the best of my ability to determine, there have been:

131 certainly cultivated, marked wild (plants in orderly rows in flower beds, landscape plantings, hanging baskets, houseplants, exotic trees not known to spread from cultivation, etc.)
90 probably cultivated, marked wild (mostly street trees and commonly cultivated plants that could conceivably be escapes or otherwise wild, but are almost certainly planted, just lacking in enough photo context clues to be certain)
1150 probably wild, marked wild (when in doubt, or totally confused, I chose this category)
131 certainly cultivated, marked cultivated
5 probably wild, marked casual (for some reason other than "captive/cultivated")
2 probably wild, marked cultivated (both were chickory, which were growing as weeds in a lawn)

...so for 1509 tracheophytes in the Blacksburg area:

76%-83% observations were of wild plants;
17%-23% observations were of cultivated plants;

of the cultivated plants:
37% were definitely incorrectly marked as wild;
62% were likely incorrectly marked as wild.
37% were definitely correctly marked as wild.

Posted on January 13, 2020 02:09 by ddennism ddennism | 4 comments | Leave a comment

October 08, 2019

Goldenrod Galls

Solidago and Euthamia are two plant genera commonly called goldenrods. They host a number of gall-making insects.

This journal entry is an attempt to catalog all the gall-making insects on goldenrods. This is probably never going to be a complete list, but I'm striving to include as many as I can find. I'm defining "gall" as a structure primarily composed of plant tissue that is induced by a resident insect and is not otherwise produced. This definition is a little wishy-washy, and could include other stem-borers and leafminers, for example, but I have excluded those creatures for now. This is not an identification guide for all insect residents of goldenrod galls. In fact, many times the current residents of a gall are not the original gall-makers! In some galls, parasitoids and inquilines are more commonly reared than the initial gall-maker.

This entry was motivated by my observation in summer 2019 that many identifications of Rhopalomyia solidaginis summer galls were misidentified on iNaturalist. I think the computer vision algorithm was partly to blame for this; it was inaccurately suggesting R. solidaginis as an identification for a great diversity of input images. These included images of normal rosettes from plants from a variety of families with no evidence of an insect or a gall being present. Users also may have been unaware of the great diversity of gall-midges and other gall-makers on goldenrods, selecting R. solidaginis by default for any leafy gall.

That said, many iNaturalist observations of galls on Solidago and Euthamia don't seem to match those of any known species of gall-maker. There may be quite a few out there to be discovered.

Similar Resources

Much of this information comes from the published work of Netta Dorchin (see full reference list at bottom).

Quick Guide to the Most Frequently-Observed Galls

Leaf Spots and Blisters

Bud Galls

  • Dasineura folliculi

  • Procecidochares atra

    • photo of Procecidochares atra bud gall on Solidago shoot apex
      initially appearing in groups along the stem, reminiscient of Brussels sprouts
      photo by Jason Dombroskie (@jasondombroskie)
      (CC BY-NC 4.0)

    • photo of clustered Procecidochares atra galls
      developing into artichoke-shaped galls, sometimes including a larger one at the apex
      often confused with R. solidaginis galls
      photo by Yann Kemper (@kemper)
      (CC0)

  • Rhopalomyia capitata

    • photo of Rhopalomyia capitata bud gall on Solidago shoot apex
      very dense cluster of short leaves with no clear organization,
      usually resulting in a flat-topped profile
      photo by Miriam Kniaz (@miriamkniaz)
      (CC BY-NC 4.0)

  • Rhopalomyia solidaginis

    • photo of Rhopalomyia solidaginis bud gall on Solidago shoot apex
      less-dense cluster of leaves of variable lengths, organized into 1-to-few rosettes,
      usually resulting in a tufted profile
      often confused with P. atra gall clusters, which are attached to the side of the stem (even if congested) and have one large, hollow cavity each (cf. the tiny, conical insect chambers of R. solidaginis, which are located in the center of each rosette, and are attached to the top of the stem, not branching from the side)
      photo by Brad Walker (@edgarallenhoopoe)
      (CC BY-NC 4.0)

"Capsules" among the Leaves and Flowers

Stem Swellings

Full Listing of Goldenrod Galls

() = uncertainty in the literature about whether this is a host species, or a note that it more rarely, conditionally, or "accidentally" serves as a host, or a species name included to account for a taxonomic change
(()) = host not mentioned in the literature, but I suspect there might be some observations of this species serving as a host on the internet.

insect host description and notes representative images sources
on Euthamia:
Asphondylia Cecidomyiidae Members of this gall midge genus develop in chambers composed of modified plant tissue with an inner fungal lining. For several species the walls of the chambers are comprised of leaf tissue that has the appearance of being glued together, forming either a rigid cone at the site of a bud or a "spot-welded" blister between two leaves. Others develop within modified flower-heads. There are often several generations per year. The organ and host species may vary throughout the year. Dorchin et al. (2015)
A. pseudorosa E. graminifolia,
((E. caroliniana))
bud-rosette,
capitula,
leaf-snap
Several different gall types per year, always on Euthamia. The most apparent ones are the bud-rosette galls formed usually on apical buds. These feature a single rosette with broadened outer leaves surrounding appressed inner leaves, the innermost much shorter, forming a single, rigid cone. This chamber is lined with white mycelium. After the insect leaves or dies, the innermost leaves turn black and wither, leaving the outer, broadened rosette. Before this happens, the gall resembles a green rose flower. Unlike in R. lobata galls, also found on Euthamia buds, spongy tissue does not form, and each bud gall contains only one inner larval chamber. A second gall type forms later in the season in developing flower head (capitula) buds. These are difficult to distinguish from normally developing capitula, though they do not flower, and instead house the developing larva. These are also lined with a white mycelium. A third type, leaf-snap galls, are pictured in Dorchin et al. (2015) but not mentioned in the text. They appear to be formed near terminal bud, and are presumably also lined with fungus. Felt (1907) raised Camptoneuromyia flavescens, another cecidomyiid, probably from these galls - specifically the flower-head ones. He attributed these galls to "Asphondylia monacha", which is a name he used to refer to what are now known as several distinct Asphondylia species. photo of Asphondylia pseurorosa gall on Euthamia lateral shoot
photo by Brad Walker (@edgarallenhoopoe)
(CC BY-NC 4.0) photo of Asphondylia pseurorosa gall on Euthamia terminal shoot
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
Dorchin et al. (2015)
Asteromyia Cecidomyiidae Members of this gall midge genus usually develop in flat blisters or leaf-spots between the faces of a single leaf. The galls are composed of discolored plant tissue with an inner fungal lining. There are often several generations per year.
A. euthamiae Euthamia sp. leaf spot
(stem spot)
Black blisters on leaves (and less-frequently on broadened, flattened stems). The blisters are lined with white mycelium. There are several generations per year. Photo of Asteromyia euthamiae galls on Euthamia leaves
photo by cassi saari (@bouteloua)
(CC BY-NC 4.0)
Photo (2) of Asteromyia euthamiae galls on Euthamia leaves
photo by Sequoia Wrens (@astrobirder)
(CC BY-NC 4.0)
Stireman et al. (2010)
Dasineura Cecidomyiidae Members of this large genus of gall midges have diverse life histories and gall forms. The species known from North American goldenrods make deformed and swollen terminal shoot buds with clustered and blistered leaves. The orange larvae develop between the leaves, but exit the gall and pupate in the soil. The shoot sometimes recovers from their presence and grows beyond the gall, leaving a deformed section of stem with very short internodes.
D. carbonaria E graminifolia bud Shoot tip bud galls, formed by several variously-adherent and contorted leaves. Circular discolored feeding spots are often visible, and these may also contribute bumps and wrinkles to the gall. The gall itself may be green to purple in color. The galls are not sealed; the larvae freely come and go to feed on the leaves, finally exiting to the soil to pupate.
A monogenous species, as is Dasineura folliculi: the offspring of a single female is all one sex, though multiple oviposition events from multiple mothers can occur on a single host bud, giving rise to mixed-sex composite galls. Because most galls house larvae of a single sex, though, individual galls may be called "male" or "female". I do not know whether male and female galls differ morphologically.
The name "carbonaria" implies a blackened structure, but this is misleading. This midge species has this name because it was mistakenly assigned to the galls made by Asteromyia carbonifera
Photo of a D. carbonaria gall on the terminal bud of Euthamia
photo by Michael K. Oliver (@cichlidmike)
(CC BY-NC 4.0)
Photo (2) of a D. carbonaria gall on the terminal bud of Euthamia
photo by Jeff Skrentny (@skrentnyjeff)
(CC BY-NC 4.0)
Dorchin et al., (2007),
Dorchin et al. (2009b)
Epiblema Tortricidae Larvae of this moth genus (at least those that form goldenrod galls) bore into stems, where they initiate narrow swelllings.
E. desertana E. graminifolia stem Very narrow stem swellings. Larvae overwinter in the gall. photo of Epiblema desertana gall on Euthamia
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
See also Miller (1963) for a reference photo (pg. 67, Fig. 3d)
Miller (1963)
Lasioptera Cecidomyiidae Members of this gall midge genus have a variety of life histories, but most form stem galls. Gagné & Jaschhof (2017)
L. cylindrigallae E. graminifolia stem Very, very narrow stem swellings. Gagné & Jaschhof (2017)
Galeopsomyia Eulophidae Chalcidoid wasps
G. haemon Asteraceae gall-within-gall This hymenopteran induces the plant to produce dark, grayish spherical structures within Asphondylia galls, each of which contains a wasp larva. Dorchin et al. (2015) found these galls most frequently within leaf snap galls, but also found them in bud galls made by A. solidaginis, A. rosulata, and A. pseudorosa. photo of Galeopsomyia haemon galls within a Asphondylia solidaginis leaf-snap gall
photo by Beatriz Moisset
(CC BY-SA 4.0)
Dorchin et al., (2015)
Rhopalomyia Cecidomyiidae Gall midges that can induce a variety of galls on a variety of hosts, but those on goldenrods tend induce the plant to form small insect chambers with soft walls, either in the context of a large, spongy mass, leaf clusters, conical structures protruding from stems and leaves, or a deformed flowering head (capitulum). Dorchin et al., (2009)
R. fusiformae E. graminifolia,
E. tenuifolia
inflorescence, mostly Very similar to the galls formed by R. pedicellata, but without a pedicel, often lacking even some of the bottom tapering section, appearing as though welded to the host tissue. This is a minor difference in gall shape, but it correlates with differences in insect morphology. context shot of many R. fusiformae galls
photo by Sequoia Wrens (@astrobirder)
CC BY-NC 4.0
Dorchin et al., (2009)
R. lobata E. graminifolia,
E. tenuifolia
bud Multi-chambered galls in apical and lateral buds. They start as 1 cm globular swellings within shoot tips or clustered around the shoot tips. Several leaves surround the spongy mass at the gall base, which grows to 6 cm. Eventually the leaves loosen and the whitish tissue reveals 5-35 larval chambers. The leaves extend beyond the gall, thinning towards the apex. Photo of R. lobata galls on lateral buds of Euthamia
photo by Jason Michael Crockwell (@berkshirenaturalist)
CC BY-NC-ND 4.0
Photo (2) of R. lobata galls on lateral buds of Euthamia
photo by Christian Grenier (@krisskinou)
CC0 1.0
Dorchin et al., (2009)
R. pedicellata E. graminifolia inflorescence, mostly Pod-like structures attached to stems, leaves, and/or inflorescences. Delicate, slender gall with a single chamber. Green to purplish-red with longitudinal ridges, tapered at both ends. Proximal end has a long, slender stalk ('pedicel') that attaches to the rest of the plant. Two generations per year, at least. Close up of R. pedicellata gall on Euthamia
photo by Sequoia Wrens (@astrobirder)
CC BY-NC 4.0
Context shot of R. pedicellata galls on Euthamia
photo by Sara Rall (@srall)
CC BY-NC 4.0
Dorchin et al., (2009)
on Solidago:
Asphondylia Cecidomyiidae Members of this gall midge genus develop in chambers composed of modified plant tissue with an inner fungal lining. For several species the walls of the chambers are comprised of leaf tissue that has the appearance of being glued together, forming either a rigid cone at the site of a bud or a "spot-welded" blister between two leaves. Others develop within modified flower-heads. There are often several generations per year. The organ and host species may vary throughout the year. Dorchin et al. (2015)
A. monacha S. juncea, S. erecta, S. uliginosa, S. altissima bud Early Spring Generation (only observed on S. altissima): Bud galls directly off of rhizomes at the soil line: Wider and harder than normal buds, single chamber lined with white mycelium. Or, slightly later in the season, bud galls at the tip of longer sprouts, stunting them and making them slightly thickened.
Summer Generation (on S. juncea, S. erecta, S. uliginosa, but not S. altissima): Much more conspicuous apical rosette bud galls, lined with mycelium, 15-30 rosette-units, forming a spherical gall complex at the shoot apex. Occasionally found on lateral buds on S. uliginosa, but rarely found there on other host species. S. uliginosa-derived adults were smaller in size as well. The authors speculated that these might represent a separate species (but distinct from the S. uliginosa-galling Asphondylia entity below).
photo of spring generation A. monacha bud gall
early spring gall
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
A. monacha bud gall cluster
summer bud gall
photo by Sequoia Wrens (@astrobirder)
(CC BY-NC 4.0)
Dorchin et al. (2015)
A. rosulata S. rugosa, S. gigantea, S. uliginosa, S. altissima leaf snap,
bud
Spring-Early Summer: Snap Galls (either hosts): Multiple leaves appear joined together at a blistering point (actually the leaves are "glued" together when the leaves are still developing) to make a single chamber lined with white mycelium. Unlike those produced by A. solidaginis, these galls are often located very near the plant apex, giving rise to a gradient from leaf snap to bud galls. This gradient is visible in the example observation.
Mid-Late Summer: Bud galls (only on S. rugosa) and only on apical buds. A single, conical chamber in the middle of a rosette of leaves. The chamber is lined with white mycelium. These galls are smaller and flatter than those formed by R. solidaginis on S. rugosa, and are composed of fewer leaves. Unlike those of R. solidaginis, these galls contain a single chamber.
Photo of a gradient of A. rosulata galls, from leaf-snap galls to a bud gall, on Solidago rugosa
bud gall and leaf-snap galls on Solidago in subsect. Venosae in Tennnessee.
photo by Ashley M Bradford (@ashley_bradford)
(CC BY-NC 4.0)
Photo of A. rosulata leaf-snap gall on Solidago rugosa
leaf-snap gall on S. rugosa in Pennsylvania
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
Dorchin et al., (2015)
A. silva S. caesia bud Very small, single-chambered bud galls at shoot tips. Several very short leaves press together to form a single, mycelium-lined chamber. photo of an A. silva gall on Solidago caesia
gall on S. caesia in Pennsylvania
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
Dorchin et al., (2015)
A. solidaginis S. altissima, S. gigantea bud Spring-Early Summer: Leaf-Snap Galls (either host): Multiple leaves appear joined together at a blistering point (actually the leaves are "glued" together while the leaves are developing) to make a single chamber lined with white mycelium.
Mid-Late Summer: Bud galls (only on S. altissima) on apical and/or axillary buds (3-5 cm in diameter), with a single, conical chamber in the middle that is lined with white mycelium. Unlike in Rhopalomyia solidaginis galls, the central chamber is not obscured by the surrounding modified leaves; it is visible without dissection. The gall walls are lined with thick white mycelia. The surrounding rosette of bunched leaves is also smaller in size, and flatter (not tufted). R. solidaginis bud galls usually contain several chambers per gall; those of A. solidaginis contain a single chamber. Another cecidomyiid fly: Camptoneuromyia adhesa sometimes emerges from snap-galls like these.
A. solidaginis leaf snap galls photo by Lena Struwe
leaf snap galls
photo by Lena Struwe (@vilseskog)
(CC BY-NC 4.0)
A. solidaginis bud gall photo by Timothy Frey
bud gall
photo by Timothy Frey (@calconey)
(CC BY-NC 4.0)
A. solidaginis leaf snap gall with pupal exuvium
leaf snap gall with protruding pupal exuvium
photo by Vitaly Charny (@vcharny)
(CC BY-NC 4.0)
Dorchin et al. (2015)
Felt (1907)
A. sp.1 (S. bicolor-galler) S. bicolor rosette A. monacha-like galls (and insects) that are distinct from A. monacha according to a molecular phylogenetic analysis. Could be the same species as A. sp. "S. sempervirens galler". One insect from a S. uliginosa rosette gall also sorted into this clade, while others from that host species sorted into A. monacha photo of a dense, A. monacha-like gall on Solidago bicolorgall on S. bicolor
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
See also Fig. 6 in Dorchin et al. (2015).
Dorchin et al., (2015)
A. sp.1 (S. sempervirens-galler) S. sempervirens bud A. monacha-like galls (and insects) that are distinct from A. monacha according to a molecular phylogenetic analysis. Could be the same species as A. sp. "S. bicolor galler". One S. uliginosa rosette gall adult also sorted into this clade, while others sorted into A. monacha. Unlike A. monacha, this species also makes lateral bud galls.
See Charley Eiseman's (@ceiseman) blog post for a photo by Noah Charney of this gall, along with details of its discovery and insects reared from it.
See also Fig. 5 in Dorchin et al. (2015) for a photo of these terminal bud galls.
Dorchin et al., (2015)
A. sp.1 (S. uliginosa-galler) S. uliginosa rosette See comments for A.  sp. (S. sempervirens galler) and A. sp. S. bicolor galler". Distinct, at least, from A. monacha, though that species also forms rosette bud galls on S. uliginosa. Dorchin et al., (2015)
A. sp.2 S. nemoralis leaf snap Leaf snap galls similar to those made by Asphondylia species are observed rarely on this species, but the agent responsible is unknown. Dorchin et al., (2015)
A. sp.2 S. tortifolia° bud A. rosulata-like galls have been observed in October, but the agent responsible is unknown. Dorchin et al., (2015)
A. sp.2 S. patula° bud Aggregated bud galls with mini-rosettes, like those made by A. monacha, have been observed on this species, but the insect remains unknown. Could be A. monacha, or another gall-maker.
My note: These observations: here and here of bud galls on S. patula may be the same galls referred to by Dorchin et al. (2015). They were found in close vicinity to typical A. monacha galls on their typical host, S. juncea.
Photo of aggregated bud galls on S. patula
Aggregated bud galls atop S. patula
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
Dorchin et al., (2015), personal observations
A. sp.2 S. odora bud Aggregated bud galls like those made by A. monacha have been observed on this species, but the insect remains unknown. Could be A. monacha, or another insect.
My note: Alvin Diamond (@adiamond) has observed these galls in Alabama on S. odora that resemble R. solidaginis bud galls on S. altissima. These may or may not be the same S. odora galls Dorchin et al. (2015) refer to.
Dorchin et al., (2015) , this observation
Asteralobia Cecidomyiidae
A. solidaginis S. pacifica bud East Asia.
Renamed Schizomyia solidaginis in Elsayed et al. (2018)
Gagné & Jaschhof (2017)
Elsayed et al. (2018)
Asteromyia Cecidomyiidae Members of this gall midge genus usually develop in flat blisters or leaf-spots between the faces of a single leaf. The galls are composed of discolored plant tissue with an inner fungal lining. There are often several generations per year.
A. carbonifera Solidago leaf spot Black blisters on leaves, lined internally with white mycelium. Interesting evolutionary biology research has been done in this system, particularly in the lab of John Stireman at Wright State University. Different lineages of A. carbonifera induce differently-shaped galls. The fungus that lines the interior of these galls is Botryosphaeria dothidea. Photo by Asteromyia carbonifera gall on Solidago leaf
photo by Matt Parr (@ginsengandsoon)
(CC BY-NC-SA 4.0)
Photo of Asteromyia carbonifera gall on Solidago leaf
photo by Sequoia Wrens (@astrobirder)
(CC BY-NC 4.0)
Photo of Asteromyia carbonifera galls on Solidago leaf
photo by Chuck Sexton (@gcwarbler)
(CC BY-NC 4.0)
Stireman et al. (2010)
bugguide
A. modesta Solidago, Erigeron, Conyza°°, Grindelia, Symphyotrichum leaf spot Leaf blisters. The larvae reside in cryptic pockets of leaf tissue that may be purple but are often nearly the same green color as the surrounding leaf tissue. The chambers are lined with a thin mycelium layer.
The species is probably polyphyletic as currently circumscribed, with two distinct clades. One clade is itself polyphyletic if A. tumifica is separated from A. modesta. Both clades include some individuals sampled from galls on Solidago. Charley Eiseman (@ceiseman) accidentally reared this midge from a leaf with more prominent leaf-mines, and photographed both the blister and the midge.
Stireman et al. (2010)
Bug Tracks post
A  tumifica Solidago stem Spongy outgrowth that partially or wholly encircles a stem. Sometimes very low on stem.
Nested within one of two A. modesta clades, rendering that clade paraphyletic. Perhaps this insect taxon will be folded into a revised concept of A. modesta in the future, or perhaps that taxon will be split up.
Photo posted to bugguide here by John van der Linden, identified by Raymond Gagné
Photo of an A. tumifica gall
photo by Tom Murray
(CC BY-NC 4.0)
Stireman et al. (2010)
Dasineura Cecidomyiidae Members of this large genus of gall midges have diverse life histories and gall forms. The species known from North American goldenrods make deformed and swollen terminal shoot buds with clustered and blistered leaves. The orange larvae develop between the leaves, but exit the gall and pupate in the soil. The shoot sometimes recovers from their presence and grows beyond the gall, leaving a deformed section of stem with very short internodes.
D. folliculi S. rugosa, S. gigantea, ((S. altissima)), ((S. canadensis)) bud Shoot tip bud galls that resemble other bud galls, but are looser and show evidence of feeding (yellowish spots, sometimes deforming the leaves somewhat) on the more-distal portions of the gall leaves. The galls may be hairy or smooth depending on the host species. The larvae exit the galls to pupate in the soil. Dasineura larvae are orange. Similar, but smaller and white-colored larvae present in galls may be Macrolabis americana, an inquiline. D. folliculi is a monogenous species, as is D. carbonaria: the offspring of a single female is all one sex, though multiple oviposition events from multiple mothers can occur on a single host bud, giving rise to mixed-sex composite galls. Because most galls house larvae of a single sex, though, individual galls may be called "male" or "female". I do not know whether male and female galls differ morphologically. Photo of a D. folliculi bud gall
on S. rugosa in Pennsylvania
photo by Daniel McClosky
(CC BY 4.0)
Dorchin et al. (2006),
Dorchin et al. (2007),
Dorchin et al. (2009b)
D. virgaureae S. virgaurea variable? Eurasia. There are several descriptions of the galls caused by this midge. Galls in shoot tips, capitula, leaf rolls, and swollen flower buds have all been ascribed to this fly. Dorchin et al. (2006)
Epiblema Tortricidae Larvae of this moth genus (at least those that form goldenrod galls) bore into stems, where they initiate narrow swelllings.
E. scudderiana S. altissima, Solidago spp., ((Heterotheca subaxillaris)) stem Narrowly cylindrical-ellipsoid stem-swelling galls. Sometimes irregularly shaped. Two holes. One small one near the top and close to a leaf axil. This is the entry hole. Another hole gets larger through the season. This is the frass-ejection and eventual exit hole. This "bung hole" is blocked with caterpillar-derived material that fits the hole closely and resembles a train wheel. Univoltine; larvae overwinter in the gall. Before winter, the caterpillar spins a silk funnel that guides the emerging moth to the exit hole. Branches often proliferate at or above the gall. But see also Lasioptera galls. photo of Epiblema scudderiana gall on Solidago stem
Photo by David (@davidenrique)
(CC BY-NC-SA 4.0)
Miller (1963),
Miller (1976),
Brown et al. (1983)
Eurosta Tephritidae Most (all except E. latifrons?) of these Tephritid fruit flies develop in bulbous galls on stems of Solidago species, usually at least partially underground (rhizomes). The most commonly observed species develops on above-ground stems.
E. comma (S. juncea, S. missouriensis, S. rugosa) rhizome Swellings on rhizomes very near soil line. Sometimes peanut-like in outline. Steyskal & Foote (1977) give a reasonable rationale for explaining that earlier authors have confused the identities of the hosts of E. elsa and E. comma; they attempt to correct the record by assigning E. elsa to S. juncea and E. comma to S. rugosa. Current databases (according to ITIS) synonymize E. elsa with E. comma. photo of Eurosta comma gall on Solidago juncea rhizome
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
bugguide, Cedar Creek (2000), Novak & Foote (1980)
E. cribata S. juncea, S. sempervirens rhizome "Crown Gall" that begins basically at the soil line (or just under), but grows upwards and is mostly above-ground at maturity. Like those of E. comma, the galls resemble peanuts somewhat. Ming (1989) included E. conspurcata and E. reticulata in synonymy with this species. My note: Are S. juncea and S. sempervirens really hosting the same fly species? photos and illustrations in Novak & Foote (1980) (paywalled) bugguide, Arthr. Fl., Sutton & Steck (2005)
E. fenestra ? rhizome Sutton & Steck (2005) say that this is also a member of the E. comma species complex. They mention that it's probably never actually been found in FL, despite earlier reports, which were due to misidentifications of E. floridensis or other members of the E. comma species complex. photos and illustrations in Novak & Foote (1980) (paywalled) Sutton & Steck (2005)
E. floridensis S. fistulosa rhizome Galls are similar to those made by E. comma and E. fenestra. Arth. Fl., Sutton & Steck (2005)
E. lateralis S. chapmanii stem Similar to the common "ball galls" made by E. solidaginis, but the gall radius is much smaller (Foster, 1934, as "E. nicholsoni", later realized to be synonymous with E. lateralis by Foote (1964)). Another synonym: E. donysa. Only known from Brevard Co., Florida, at least recently. It may be critically endangered (Sutton & Steck (2005)), or even extinct. They give S. odora as the host, but the host is presumably S. chapmanii based on location, which was not regularly segregated from S. odora. They point out a very old record by Wiedemann (1830) also possibly of this species in the "Indien" (sic) River area of Florida. Foster (1934) points to galls found "near Titusville", "near Malabar", and "from 5.5 miles southwest of Indian River" all near the coast. Arth. Fl., Sutton & Steck (2005)
E. solidaginis S. altissima, S. gigantea, (S. canadensis), (S. rugosa) stem Nearly spherical galls, made on the aboveground stem, rather than on the rhizome like most of the rest of the goldenrod-galling members of this genus. The exterior vestiture of the gall depends on the identity of the host species. They are hairy when on S. altissima (presumably also when on S. canadensis and S. rugosa), but smooth and shiny when on S. gigantea. There is a great wealth of literature on the evolutionary dynamics of this system. Gall diameter seems to be under the control of the insect's genetics, not the host plant's. Insects that produce galls with larger diameters are more likely to survive attack by parasitic wasps, whose ovipositors are unable to penetrate the thicker galls. However, larger galls are more attractive to birds, which eat the larvae in winter. There is also interesting research on host-species specialization by different populations of this fly (on S. altissima vs. on S. gigantea), and the divergent selective pressures at play. The galls are so frequent on S. altissima in the mid-Atlantic that the presence of galls has been suggested as an identification aide for distinguishing S. altissima from S. canadensis, although S. canadensis can also be (more rarely?) a host for this fly. photo of E. solidaginis stem gall on Solidago altissima
on S. altissima
photo by Daniel McClosky
(CC BY 4.0)
photo of E. solidaginis stem gall on Solidago gigantea
on S. gigantea
photo by Daniel George (@danielgeorge)
(CC BY-NC 4.0)
Bucknell University Solidago Gall Website, Moffatt et al. (2019), Stoltzfus (1989)
Eutreta Tephritidae Tephritid fruit flies whose larvae bore through stems, usually inducing galls. They have a variety of hosts in Asteraceae and Verbenaceae.
Eutreta hespera Solidago sp. rhizome Reared once from rhizomes of "a goldenrod" near Custer, South Dakota, but the adult flies have been collected along streams and grassy slopes from the Dakotas westward, through much of western North America. Stoltzfus (1974)
Eutreta novaeboracensis Solidago rugosa, (Solidago spp.) rhizome, stem Larvae bore through both rhizomes and above-ground stems, inducing galls as swellings of those organs. The stem-borers emerge earlier than the rhizome-borers (and are bivoltine rather than univoltine), so these two groups might represent cryptic sister species. Stem-galls can be found near the ground, sometimes described as crown galls. "Eutreta sparsa" is sometimes attributed to these and other galls on North American Astereae, but this is actually a South American species that probably does not make galls on Solidago, instead associating with Stachytarpheta (in Verbenaceae) branches. drawing of Eutreta novaeboracensis galls on the rhizomes of Solidago altissima
drawing by Millett T. Thompson (1907)
(public domain)
bugguide, Stoltzfus (1974), Thompson (1907)
Galeopsomyia Eulophidae Chalcidoid wasps
G. haemon Asteraceae gall-within-gall This hymenopteran induces the plant to produce dark, grayish spherical structures within Asphondylia galls, each of which contains a wasp larva. Dorchin et al. (2015) found these galls most frequently within leaf snap galls, but also found them in bud galls made by A. solidaginis, A. rosulata, and A. pseudorosa. photo of Galeopsomyia haemon galls within a Asphondylia solidaginis leaf-snap gall
photo by Beatriz Moisset
(CC BY-SA 4.0)
Dorchin et al., (2015)
Gnorimoschema Gelechiidae Larvae of some members of this large moth genus develop within stem swellings on goldenrods. Several species have been described from goldenrods, but not all are universally accepted as distinct. The larvae bore into the shoot apex, traveling down the center of the stem while feeding and molting. Deformed, stubbier leaves at the shoot apex give a clue of the presence of the larva before it begins to make its gall. The larvae then backtrack upwards, where they induce a stem swelling with a large internal cavity. There is one larva per gall. Before spinning a cocoon they bore an adult exit hole. This hole is variably capped with plant- and/or insect-derived substances. Miller (2000) gave evidence that these "bung hole" structures were diagnostic of particular species of moth, arguing that differences in their construction were not merely consequences of different host goldenrod building material, but were intrinsic to the species of insect. Whether all the species in his monograph on the group are distinct enough to warrant their own names (particularly in the case of G. jocelynae) remains debated, but with some support from molecular phylogenies (Nason et al., 2002). Miller gave species names with neuter suffixes, but I don't know which form is correct according to nomenclatural rules so I follow the suffixes present in online checklists and more recent literature instead.
G. gallaeasteriella S. flexicaulis, Symphyotrichum saggitifolium, (Solidago caesia, S. uliginosa, Eurybia divaricata, and perhaps many others) stem No bung is formed; the adult exit hole is difficult to see because it is capped with plant epidermal tissue instead. It is located in the top third of the gall, like most other goldenrod Gnorimoschema galls, but unlike those of the otherwise very similar G. gallaespeciosum, which are instead formed near the gall equator.
This species might be conspecific with G. gallaediplopappum, a moth with unknown gall biology (Miller).
Unlike other Gnorimoschema species, the leaves at the shoot apex may not be deformed by the presence of the larva, but the shoot itself may have its growth arrested.
Are all these different host reports really referring to the same species of moth? They are all basically woodland Astereae, but I can't help but wonder whether there have been some mistakes in host identification. Miller (2000) reported that he could only find these galls on Symphyotrichum saggitifolium, from which he reared adults that matched Kellicot's (the original species authority). Judd (1962) could not find any galls on S. caesia despite there being large numbers of these plants adjacent to and within a S. flexicaulis site with many galls. I think it's likely that the initial report of galls from S. caesia may be in error since these two Solidago species were sometimes confused or lumped early on. Then again, Eurybia and Doellingeria (??) host reports suggest a wider range of acceptable hosts. Busck (1911) considered this problem of host identification, and concluded that the species may simply have a wide host range. His "S. latifolia" is our S. flexicaulis; his Aster corymbosus apparently refers to E. divaricata. I still can't help but wonder whether all these hosts might really be misidentified plants in the Symphyotrichium cordifolium species complex (which includes S. saggitifolium), which are variable and difficult to identify in the absence of flowers.
woodcut of gall by D. S. Kellicott (1878)
woodcut of gall by D. S. Kellicott (1878)
Gnorimoschema gall on Solidago caesia
Gnorimoschema gall on S. caesia
photo by Rob Curtis (@rcurtis)
(CC BY-NC-SA 4.0)
Nazari & Landry (2012), Miller (2000), Judd (1962), Kellicott (1878), Busck (1911)
G. gallaesolidaginis S. altissima, S. canadensis, (S. gigantea) stem Probably the most common Gnorimoschema moth that forms galls on goldenrods. These are ellipsoid stem galls with an exit hole stuffed with characteristically light-colored "bung". The bung tissue is essentially flush with the surrounding plant tissue, which often forms a slightly raised ring around the hole. The galls are wider than those made by Epiblema moths. Miller (1963) mentioned that there are other species in this genus that make galls on other Solidago species. His later monograph on them (2000) listed eight species, most of which have been reported from Solidago species. Later, Heard & Kitts (2012) compared G. gallaesolidaginis on S. altissima and S. gigantea. Nason et al. (2002) considered this to either contain a single differentiating species (into semispecies) onto the two respective host-groups (S. altissima / canadensis and S. gigantea), or two barely-isolated cryptic species, in which case G. jocelynae is the species name for the group that feeds on S. gigantea. photo of Gnorimoschema gallaesolidaginis spindle gall on Solidago stem
photo by Colin D. Jones (@colindjones)
(CC BY-NC 4.0)
photo of empty Gnorimoschema gallaesolidaginis spindle gall on Solidago altissima stem
photo by Daniel McClosky (@ddennism)
(CC BY4.0)
Miller (1963), Heard & Kitts (2012), Nazari & Landry (2012)
G. gallaespeciosum (S. speciosa, S. jejunifolia, S. rigidiuscula, S. pallida) stem Elliptical stem galls similar to those of G. gallaesolidaginis, but differing in exit hole placement nearer the equator and the lack of bung tissue (instead being capped with plant epidermis). Reported from S. speciosa, but the type specimen is from Ramsey Co., MN, which is out-of-range for S. speciosa in the new, strict sense. The host is more likely S. jejunifolia and/or S. rigidiuscula, goldenrod species segregated from S. speciosa in the broad sense that are native to Ramsey County. S. pallida was also segregated from S. speciosa, so it is also included here as a potential host. Miller (2000)
G. gibsoniella S. rigida, Symphyotrichum pilosum stem S. rigida was reported as the host plant for the type collection, but Miller (2000) reared these from galls on Symphyotrichum pilosum stems. The initial description by Busck here notes that it forms galls just above the ground, and Miller also noted this, mentioning that he only found them after mowing tall grass. Nazari & Landry (2012), Busck (1915)
G. jocelynae S. gigantea stem Very similar to galls by G. gallaesolidaginis, but with dark bungs that are recessed into the exit hole. The bung material is not flush with the surrounding plant epidermis. This is the name Miller (2000) gave to the host-race derived from G. gallaesolidaginis, when that species established a cryptic sister species on S. gigantea (Nason et al. 2002). The main difference is in the bung hole coloration - which was determined by the adult's ancestral host species, not its current host species in Miller's larva transplant experiments, however, Nason et al. (2002) points out that these exit hole characteristics could be idiosyncratic to particular plants and Miller only examined three specimens for this experiment. I wonder whether the larva transplant experiments might have a different interpretation than Miller's: The bung material could reflect the host species that the larva spent most of its life feeding on, which in his experiments would be the plant with the donor-gall, not the plant with the receiver gall. Works after Nason et al. (2002) refer to these moths as "G. gallaesolidaginis, gigantea host-race". Nazari & Landry (2012)
G. salinaris S. sempervirens, S. missouriensis, S. juncea, and perhaps other members in this species group; (S. gigantea, S. ulmifolia) stem Differences from G. gallaesolidaginis galls: (1) bung material is dark brown to black in color, (2) galls tend to be lower on the stem, (3) the interior surface of the bung (not visible from outside) is cushioned by layers of spongy silk material, (4) different host species. The original description, by Busck, noted that the insects were reared from galls similar to those of G. gallaesolidaginis, but on S. sempervirens. Miller (2000) associated this species with additional hosts inland: the S. juncea species complex. He referred to these plants as "S. juncea/missouriensis", but based on the locations of his collections, he was almost certainly only collecting galls from S. juncea, although S. missouriensis sporadically occurs that far east. Nazari & Landry (2012) list S. gigantea as a host according to one Michigan record, but this might be G. gallaesolidaginis/G. jocelynae. Miller (2000) attributed some galls on S. ulmifolia to this species, but noted that these were rare and only found at one site with abundant galls on S. juncea. Miller may be following a taxonomic treatment of these three main host species that does not match the current circumscriptions of these species, because some of his statements, particularly that S. sempervirens and S. juncea hybridize, sound a little strange, and he seems to imply that these three species are closely related. Both molecular- and morphology-informed phylogenies imply otherwise. I wonder whether some of Miller's hosts might have been S. uliginosa or other species. However, all reported hosts do have smooth stems in common.
Miller (2000) noted that some of the adult characteristics that Busck said differentiated this species from G. gallaesolidaginis were not as consistent as Busck claimed. He instead focused more on the gall characters (host plant ID and bung characters), and presented evidence that this species makes galls closer to the ground, and with a more variable shape in outline, ranging from elliptical to pear-shaped. The gall shape begins perfectly vertically-symmetric in the ultimately pear-shaped galls, becoming asymmetic only with age. The bung itself is dark in coloration, like that of his G. jocelynae and surrounded by a ring of tissue that may be slightly raised. The bung tissue is flush with this ring, like in G. gallaesolidaginis and unlike in G. jocelynae.
photo of Gnorimoschema spindle gall on Solidago juncea stem
Gnorimoschema gall on S. juncea
photo by Daniel McClosky (@ddennsim)
(CC BY 4.0)
Nazari & Landry (2012), Miller (2000), Busck (1911)
Janetiella Cecidomyiidae I can't find much on this one. Gagné & Jaschhof (2017) call the genus a "diverse assemblage".
J. inquilina Solidago sp. ? aka Oligotrophus inquilinus Felt 1908; on "S. canadensis", which, at the time, could have referred to several species in Solidago subsection Triplinerviae. Gagné & Jaschhof (2017)
Lasioptera Cecidomyiidae Members of this gall midge genus have a variety of life histories, but most form stem galls. Gagné & Jaschhof (2017)
L. solidaginis Solidago stem Irregular, elongated stem-swelling galls. "Knotty" in appearance according to Felt (but so many of the galls are mismatched in that work). bugguide, Gagné & Jaschhof (2017)
Lestodiplosis Cecidomyiidae Larvae of this midge genus are usually predators of other Cecidomyiid larvae, but some have been reported to be gall formers, maybe in error. Gagné & Jaschhof (2017) give the following species from Solidago galls, but are not claiming that they initiated the gall. Gagné & Jaschhof (2017)
L. carolinae S. canadensis (presumably sensu lato) bud Reported by Felt to be the initiator of the gall, but given the life history of others in the genus, more likely a predatory species? Found in a rosette bud gall on "S. canadensis". Maybe this is a synonym of Rhopalomyia carolina(e), itself a synonym of R. solidaginis? Found in Asheville, NC Gagné & Jaschhof (2017)
L. rugosae Solidago sp. New York Gagné & Jaschhof (2017)
L. triangularis Solidago sp. leaf New York Gagné & Jaschhof (2017)
Procecidochares Tephritidae
P. anthracina S. californica bud Bud galls cluster on stems near where they emerge from rhizomes. Usually buried in humus, but not truly subterranean. Univoltine, unlike P. atra. Reported from "S. velutina" in California - these plants are now in the segregate species, S. californica, according to John Semple's website. Goeden & Teerink (1997)
P. atra S. altissima, S. gigantea, S. rugosa, S. nemoralis, (Erigeron canadensis), (Aster°° sp.) bud Spring Generation: Large stem galls at the base of the host plant, each containing several larvae.
Summer Generation: Lateral bud galls that look like Brussels sprouts initially, and eventually open as the fly matures. The terminal bud is also sometimes galled, but usually in addition to lateral galls. (My observation: When the terminal bud is galled, it is often much larger than the accompanying lateral galls.) The gall chamber is large and not sealed, much larger than the fly larva or pupa, and is slightly open at the distal end. The chamber has the appearance of being inset into the stem somewhat, although the surrounding tissue may not technically be derived from the stem, but rather from other plant tissues. At maturity, the rosette of leaves surrounding the gall typically flatten and grow away from the gall, giving the gall a more rosette-like appearance.
(My note: At this stage, these galls, particularly the terminal ones, can superficially resemble those of Rhopalomyia solidaginis and Asphondylia solidaginis. However, those midge-induced galls have distinct chambers formed by one or few young leaves. The midge gall chambers are cryptic, translucent conical structures set atop the host stem rather than appearing hollowed-out within stem-like tissue.) Each gall has only one larva, unlike in the spring generation.
This species probably has many other hosts, including some outside Solidago, although some researchers have speculated that there may be cryptic host-races within P. atra, some of which may have fully speciated (Philips & Smith 1998).
photo of spring Procecidochares atra gall
photo of spring Procecidochares atra gall, cut open
Spring Generation gall
photos by Daniel McClosky (@ddennism)
(CC BY 4.0)
photo of Procecidochares atra galls
Summer Generation galls
photo by Jason Dombroskie (@jasondombroskie)
(CC BY-NC 4.0)
another photo of Procecidochares atra galls
photo by Yann Kemper (@kemper)
(CC0)
vertical section  of Procecidochares atra gall
summer generation, vertical section through terminal bud gall
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
Wikipedia, iNat obs, bugguide, Philips & Smith (1998), Aldrich (1929) for S. nemoralis record
P. minuta (Solidago), Astereae stem Recorded from a stem gall on Solidago californica in Wasbauer (1972): "C. D. A. 1 In stem gall; CALIFORNIA: Palomar Mt., San Diego Co., IX-19-1964, E. D. Algert". This species is known to produce galls on a number of composite host species, including rabbitbrushes. I'm not sure this record is from a correctly identified host plant; rabbitbrushes can look vaguely like goldenrods before blooming. Wasbauer (1972)
P. polita (S. virgata, S. chrysopsis), (((S. sempervirens))),(((S. mexicana))) ? Reared from Solidago sp. "small, roundish galls" by Girault (1913) in Virginia. Reported from galls of Solidago sp. by Johnson (1910). However, Aldrich (1929) wrote that accounts of this species being reared from Solidago galls are in error, and actually refer to P. atra due to some nomenclatural confusion at the time in the literature.
Much later, Ibrahim (1980) attributed "Solidago stricta" stem galls collected in Dade County, Florida to P. polita. At that time, "Solidago stricta" would have referred to what is now known as S. virgata or possibly S. chrysopsis at that location (see John Semple's website for details). There are older records for collections in the Jacksonville area (Sutton & Steck 2005) and the Falls Church, Virginia area (Aldrich 1929). Sutton & Steck (2005) caution that many details in Ibrahim (1980) are inaccurate and repeat known mistakes from earlier literature, though they don't mention the P. polita record specifically.
The adult flies are apparently easily distinguished from P. atra by having entirely yellow legs rather than having black femora and coxae. Goeden & Norrbom (2001) say it's distributed along the east coast, from Massachusetts to Florida. None of these post-1929 sources describe the gall.
This fly seems to be restricted to the east coast of the USA, so its host plant, if it is a Solidago species, is probably a coastal species. Wasbauer (1972) includes some records from "S. stricta" galls as well.
Aldrich (1929), Ibrahim (1980), Goeden & Norrbom (2001), Sutton & Steck (2005)
Rhopalomyia Cecidomyiidae Gall midges that can induce a variety of galls on a variety of hosts, but those on goldenrods tend induce the plant to form small insect chambers with soft walls, either in the context of a large, spongy mass, leaf clusters, conical structures protruding from stems and leaves, or a deformed flowering head (capitulum). Dorchin et al. (2009)
R. anthophila S. altissima capitulum Capitulum (flower-head) galls among the flowers of the host. Cylindrical, or like a truncated cone. Fuzzy and whitish. Inner chamber conical, resembling the shape of other Rhopalomyia insect chambers, with thin walls. Photo of an isolated R. anthophila gall on the inflorescence of Solidago altissima
photo by Kevin Keegan (@kevinliam)
(CC0)
context shot of many R. anthophila galls
among S. altissima capitula
photo by Daniel McClosky
(CC BY 4.0)
macro image of a R. anthophila gall
photo by Dan Mullen
(CC BY-NC-ND 2.0)
Dorchin et al. (2009)
R. bulbula S. juncea rhizome bud Only a spring generation is known, but the insect is presumably multivoltine;
Spring Generation: "Clustered on rhizomes, at the bases of spring shoots. The gall resembles a bud, with acute apex and base. Surface is smooth and white, with green stripes where exposed to light." Single chambered.
photograph of R. bulbula gall from Felt (1917)
photo by E. P. Felt (1917)
Dorchin et al. (2009), Felt (1917)
R. capitata S. gigantea, S. leavenworthii, (S. altissima), ((S. canadensis)) bud Spring Generation: Few (1-8) conical chambers surrounded by disorganized small leaves, sheathed (initially at least, sometimes loosening) by several wide leaves. Distinctly more conspicuous than R. solidaginis spring galls.
Summer Generation: Apical bud gall with many small leaves of uniform length in the middle, surrounding many (6-20) closed larval chambers. Wide leaves also sheath these galls. The uniformly-small leaves give the overall gall complex a flat-topped appearance. Whereas tufts of leaves that comprise the summer generation gall complex formed by R. solidaginis form discernible mini-rosettes, each surrounding a larval chamber, in R. capitata the gall leaves are not obviously so-organized, perhaps as a consequence of there being many more chambers.
My note: In the upper Great Lakes region there are leafy galls on S. gigantea that more closely resemble those made by R. solidaginis.
photo of R. capitata spring generation bud gall
spring generation
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
photo of typical summer generation R. capitata bud gall
summer generation
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
vertical cross section of typical summer generation R. capitata bud gall
summer generation bud gall, vertically sectioned
photo by hallm (@hallm)
(CC BY-NC 4.0)
R. capitata bud gall in MN
summer generation bud gall, from above, in MN
photo by Miriam Kniaz (@miriamkniaz)
(CC BY-NC 4.0)
Dorchin et al. (2009)
R. clarkei S. rugosa, S. altissima leaf outgrowth Small, conical, single-chambered. Usually on abaxial leaf surface, but can also appear on adaxial surface or on stems. When on leaves, attached at a major vein. Green to yellow-green and covered with hairs. Very young galls with a tuft of hair at base. Multivoltine. Less frequent on S. altissima.
(My observation: There are several gall observations on iNaturalist that fit this description, and are currently identified as R. clarkei, but they do not all closely resemble one another. They may represent different stages of development, or else different presentations on different host species.)
Photo of a R. clarkei leaf gall
photo by Will Van Hemessen (@wdvanhem)
(CC BY-NC 4.0)
Photo of a R. clarkei leaf gall
photo by Sara Scharf (@scharf)
(CC BY-NC 4.0)
Dorchin et al. (2009)
R. cruziana *(S. spathulata), (S. velutina) capitulum? From an unknown gall from an unidentified Solidago species growing in the Santa Cruz mountains in California. The species to the left are my speculation based on the Solidago species native to this region. Dorchin et al. (2009) infers that the gall is probably a capitulum gall because the adult insects closely resemble other capitulum-gallers in this genus. Dorchin et al. (2009)
R. gina S. juncea leaf outgrowth Like R. clarkei galls, but usually on upper side of leaf and with a corresponding scar or little tail on the opposite side. Hairless, probably reflecting the vestiture of the host plant. See Figs. 68-69 in Dorchin et al. (2009) for images.
My note: Fig. 69 shows a leaf with what might be pubescence on the abaxial surface (in addition to the normal cilia at the leaf margin), which makes me wonder whether this is really S. juncea
Dorchin et al. (2009)
R. guttata S. bicolor capitulum Forms teardrop shaped galls that retain the capitulum-pedicel. Difficult to find among regular flower-heads. Dorchin et al. (2009)
R. hirtipes S. juncea bud Forms fleshy bud galls at the shoot apex, often arresting shoot growth while the plant is still very short. Gall initially has a tapered tip, but this disappears with growth. The whole gall eventually becomes ovoid and reminiscent of a potato. Spongy and usually multi-chambered. See also R. thompsoni for a similar gall that appears earlier in the season and mostly underground. Photo of a R. hirtipes bud gall
photo by Catherine Klatt (@catherineklatt)
(CC BY-NC 4.0)
R. hirtipes bud gall photo showing the plant flowering from lateral buds under the gall
not always arresting growth
photo by Charles and Kathy Appell (@charleshappell)
(CC BY-NC 4.0)
Dorchin et al. (2009)
R. inquisitor S. gigantea leaf outgrowth? Originally described as an inquiline in R. capitata galls, but this could not be replicated by Dorchin et al. (2009). The did notice R. clarkei-like galls (except smooth-surfaced) on S. gigantea, though, particularly on leaves from within Dasineura folliculi galls, and conjectured that these R. clarkei-like galls might be the real galls occupied by this species. Perhaps Felt (original description) confused D. folliculi galls with R. capitata galls, and then concluded that R. inquisitor was an "inquiline" that way? However, Dorchin et al. (2009) were unable to rear any adults from these R. clarkei-like galls on S. gigantea, so the galls where R. inquisitor resides remain unclear. An example of these galls is probably shown here: by Sara Rall (@srall). Close up of Rhopalomyia galls on Solidago gigantea leaves
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
Photo of R. inquisitor galls, maybe, on S. gigantea leaves
click to zoom to see small leaf-outgrowths. Leaf clustering may be caused by D. folliculi. Host plant is S. gigantea.
photo by Sara Rall (@srall)
(CC BY-NC 4.0)
See also Figs. 66-67 in Dorchin et al. (2009) for reference images.
Dorchin et al. (2009)
R. racemicola S. altissima, (S. fistulosa) capitulum Green, bristly, onion-shaped capitulum galls, sometimes found in aggregations. Galls on S. fistulosa were tentatively identified as this species by Raymond Gagné . The illustration in Felt (1915) by L. H. Joutel looks suspiciously like Schizomyia racemicola, but I can find no other published images of these galls. Illustration of a R. racemicola bud galls
Illustration by L. H. Joutel in Felt (1915)
Dorchin et al. (2009)
R. solidaginis S. altissima, S. canadensis, S. rugosa, ((S. odora)) bud Spring Generation: Inconspicuous, shoot tip rosette bud-galls, often stunting the shoot. Unlike the later generation, these typically enclose a single, white, conical gall-chamber, but sometimes several gall-chambers are connected longitudinally.
Summer generation: Each of multiple (2-5) chambers is surrounded by a group of very short and narrow leaves, which are in turn surrounded by longer and wider leaves to form a distinct rosette-subunit within the gall complex. The whole complex itself forms a conspicuous mass of leaves. This is a very common gall in the mid-Atlantic states.
Alvin Diamond (@adiamond) has observed similar galls on S. odora in Alabama; I'm not sure whether these are the galls that Dorchin (2015) refers to and suspects an Asphondylia midge of making (see*Asphondylia* entries above) or if this is a distinct entity.
photo of R. solidaginis typical summer gall on Solidago altissima
summer generation
photo by Daniel McClosky (@ddennism)
(CC BY-NC 4.0)
vertical section through R. solidaginis summer generation bud gall on Solidago altissima
summer generation, vertical section
photo by Daniel McClosky (@ddennism)
(CC BY-NC 4.0)
See also Figs. 70, 72 in Dorchin et al. (2009) for images of spring generation galls.
Dorchin et al. (2009)
R. thompsoni S. altissima, (S. juncea), (S. rugosa) rhizome bud Spring Generation: Solitary or clustered, bulbous, fleshy masses with 1-8 chambers each. Start on rhizomes (underground) but become apparent above ground by emergence time in early May.
Second Generation: Brownish, globular multi-chambered swellings of the rhizomes that stay underground until late September when they become apparent above the soil surface for adult emergence. Both galls and adults resemble R. hirtipes. Dorchin et al. (2009) could only find galls that reared adults similar to the type of R. thompsoni from galls from S. atissima, but Felt had listed the other two species as hosts.
Dorchin et al. (2009)
R. sp. (S. fistulosa-stem-galler) S. fistulosa stem Aggregated stem galls, each gall a hairy grayish oval, with a single chamber each. The aggregate commonly has a star-like structure. They appear most similar to R. racemicola galls. Might be responsible for the gall in this observation. Dorchin et al. (2009)
Schizomyia Cecidomyiidae Gagné & Jaschhof (2017)
S. racemicola Solidago capitulum Greenish-purplish onion-shaped capitulum galls alongside normal capitula in the inflorescence. Gall exterior is smooth. Gall-maker larva is bright red-orange. It exits the gall as a larva and pupates elsewhere. photo of S. racemicola gall on S. ulmifolia
on S. ulmifolia
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
photo of S. racemicola gall on S. altissima
on S. altissima
photo by Daniel McClosky (@ddennism)
(CC BY 4.0)
bugguide
S. solidaginis S pacifica See Asteralobia solidaginis entry. bugguide
Tephritis Tephritidae
T. webbi Solidago sp. capitulum "M. F. Canova states that the specimen was taken from a gall in the flowerhead of goldenrod." Sycan Glen, OR. The adult insects closely resemble T. michiganiensis and T. pura, neither of which have known host species (at least by 1951). Quisenberry (1951)

There are some records of Trupanea spp. infesting goldenrod flower-heads, but do they form galls?

Wasbauer (1973) gives a secondary record for galls of Aciurina bigeloviae on Solidago, but this species probably doesn't regularly gall Solidago (?)

Foote & Blanc (1963) ascribe a collection of galls on Solidago in Inyo Co., CA to A. ferruginea, but this fly typically galls rabbit-brushes. Maybe a mistaken host ID?

See also Aster Yellows phytoplasma, which can induce phyllody in Solidago.

°observed in Maryland (??)
°°I think this Conyza species is probably C. canadensis, which is back in Erigeron now.
°°°probably refers to Symphyotrichum species now, not Aster sensu lato.

1These entries are for insects that induced galls that resembled those of A. monacha, but were found on other host Solidago species, and were divergent phylogenetically.
2These entries are for the un-studied (to my knowledge) insects that induce Asphondylia-like galls on other host Solidago species. These are known only from the appearance of galls on these goldenrods; the midges themselves have been neither reared nor described. These entries may represent infestation by known Asphondylia midges, A. monacha and A. solidaginis in particular, on occasional/accidental host species. I myself have observed Asphondylia-like galls on S. patula. They may also represent unrelated gall-making organisms.

Unexpected / Interesting goldenrod gall observations

Rhopalomyia solidaginis-like bud gall on Solidago chapmanii: https://www.inaturalist.org/observations/37226412

Asphondylia monacha-like bud gall on Solidago leavenworthii:
https://www.inaturalist.org/observations/36869277

bud-gall on Solidago sp. (?) in Kansas:
https://www.inaturalist.org/observations/34869212

Asphondylia monacha-like bud galls on Solidago sempervirens:
https://www.inaturalist.org/observations/35173501
https://www.inaturalist.org/observations/16081294
(probably the same bud-galler discovered by Charley Eiseman here:
https://bugtracks.wordpress.com/2012/08/09/goldenrod-rosette-galls-part-2/
and in the above table as: "Asphondylia sp. (S. sempervirens-galler)"

bud-gall on Solidago juncea (?):
https://www.inaturalist.org/observations/35173440

tiny bud-galls on Solidago ulmifolia (?) in IL:
https://www.inaturalist.org/observations/33584564
https://www.inaturalist.org/observations/33623087

Asphondylia-like bud galls on southeastern USA Solidago sp., possibly A. monacha on S. erecta:
https://www.inaturalist.org/observations/33794356
https://www.inaturalist.org/observations/34156586
https://www.inaturalist.org/observations/17063696
https://www.inaturalist.org/observations/31258368
https://www.inaturalist.org/observations/31420378

Asphondylia monacha-like bud gall cluster on Solidago sp. in VA:
https://www.inaturalist.org/observations/35246250

Asphondylia monacha-like bud gall cluster on Solidago altissima (?) in NJ:
https://www.inaturalist.org/observations/34508254
...or very, very large R. solidaginis galls, I guess. More-typical looking examples of R. solidaginis in immediate vicinity: https://www.inaturalist.org/observations/34507741
Could also be R. capitata but on an atypical host species (host is definitely not S. gigantea), and the gall is not flat-topped.
A similar situation in N IL:
https://www.inaturalist.org/observations/28883789

Asphondylia solidaginis-like leaf deformities (and maybe a snap-gall?) on Solidago sp. in MS:
https://www.inaturalist.org/observations/12472658

A very tightly-wound Asphondylia solidaginis-like gall:
https://www.inaturalist.org/observations/14392656

Is this Asphondylia rosulata on Solidago rugosa?
https://www.inaturalist.org/observations/29881623

not Procecidochares atra, looks kind of like Dasineura, but lateral galls present in addition to terminal gall, on S. altissima in OH:
https://www.inaturalist.org/observations/32848998

Dasineura-like leaf deformities in spring in TX:
https://www.inaturalist.org/observations/24096959

A strange bud gall on Solidago in CT:
https://www.inaturalist.org/observations/33925735

A bud gall on Solidago way out-of-range in Washington State:
https://www.inaturalist.org/observations/13619060

Rhopalomyia anthophila-like gall chamber, but not on the inflorescence, in MI:
https://www.inaturalist.org/observations/15411274

Rhopalomyia solidaginis-like bud galls in appearance, but on S. gigantea in MN:
https://www.inaturalist.org/observations/17081514
https://www.inaturalist.org/observations/29258718
and in MA:
https://www.inaturalist.org/observations/31895692

Asphondylia-like bud galls on S. rigida (?) in IL:
https://www.inaturalist.org/observations/18784886
https://www.inaturalist.org/observations/32639920

Asphondylia-like bud galls on S. buckleyi (?) in IL:
https://www.inaturalist.org/observations/60788196

bud gall on Solidago ptarmicoides in Ontario:
https://www.inaturalist.org/observations/19331736

Rhopalomyia capitata on a Solidago species that is not S. gigantea:
https://www.inaturalist.org/observations/31247515




References


Aldrich (1929):
https://repository.si.edu/bitstream/handle/10088/15820/1/USNMP-76_2799_1929.pdf

Arth. Fl. = Arthropods of Florida website by Florida State Museum of Entomology:
http://www.fsca-dpi.org/Diptera/Families/Tephritidae/Species/Eurosta/eurosta_cribrata.htm

Bucknell University Eurosta biology page:
http://www.projects.bucknell.edu/solidago/main.html

Cedar Creek Ecosystem Science Reserve (2000):
http://cedarcreek.umn.edu/insects/029061n.html

Dorchin et al. (2006):
Dorchin, N., Scott, E. R., Abrahamson, W. G. (2006) First Record of Macrolabis (Diptera: Cecidomyiidae) in America: A new inquiline species from Dasineura folliculi galls on goldenrods. Systematics 99(4): 656-661.

Dorchin et al. (2007):
Netta Dorchin, Carolyn E. Clarkin, Eric R. Scott, Michael P. Luongo, Warren G. Abrahamson, Taxonomy, Life History, and Population Sex Ratios of North American Dasineura (Diptera: Cecidomyiidae) on Goldenrods (Asteraceae), Annals of the Entomological Society of America, Volume 100, Issue 4, 1 July 2007, Pages 539–548, https://doi.org/10.1603/0013-8746(2007)100[539:TLHAPS]2.0.CO;2

Dorchin et al. (2009):
https://www.mapress.com/j/zt/article/view/zootaxa.2152.1.1

Dorchin et al. (2009b):
Dorchin, N., Scott, E. R., Clarkin, C. E., Luongo, M. P., Jordan, S. and Abrahamson, W. G. (2009) Behavioural, ecological and genetic evidence confirm the occurrence of host‐associated differentiation in goldenrod gall‐midges. Journal of Evolutionary Biology, 22: 729-739. doi:10.1111/j.1420-9101.2009.01696.x

Dorchin et al. (2015):
Dorchin, N., Joy, J. B., Hilke, L. K., Wise, M. J., Abrahamson, W. G. (2015) Taxonomy and phylogeny of the Asphondylia species (Diptera: Cecidomyiidae) of North American goldenrods: challenging morphology, complex host associations,
and cryptic speciation. Zoological Journal of the Linnean Society, 174: 265-304. doi:10.1111/zoj.12234
https://academic.oup.com/zoolinnean/article/174/2/265/2449766

Elsayed et al. (2018):
Elsayed, A. K., Yukawa, J., Tokuda, M. (2018) A taxonomic revision and molecular phylogeny of the eastern Palearctic species of the genera Schizomyia Kieffer and Asteralobia Kovalev (Diptera, Cecidomyiidae, Asphondyliini), with descriptions of five new species of Schizomyia from Japan. Zookeys 808: 123-160.

Heard & Kitts (2012):
http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.726.1531&rep=rep1&type=pdf

Ibrahim (1980):
https://ufdcimages.uflib.ufl.edu/AA/00/02/97/95/00001/AA00029795_00001.pdf

Felt, E. P. (1915):
http://www.nysm.nysed.gov/staff-publications/29th-report-state-entomologist-injurious-and-other-ins

Felt, E. P. (1917):
Felt, E.P. Key to American Insect Galls. New York State Museum Bulletin 200.
https://www.biodiversitylibrary.org/item/35191

Foote (1964):
Foote, R. H. (1964) A new synonym in the genus Eurosta (Diptera: Tephritidae). Proceedings of the Entomological Society of Washington 66 (1): 61.

Foote, R. H., Blanc, F. L., and Norrbom, A. L. (1993) Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico

Goeden & Teerink (1997):
https://www.biodiversitylibrary.org/part/55666#/summary

Goeden & Norrbom (2001) Life history and description of adults and immature stages of Procecidochares blanci, n. sp. (Diptera: Tephritidae) on Isocoma acradenia (E. Greene) E. Greene (Asteraceae) in Southern California. Proceedings of the Entomological Society of Washington 103 (3-4): 517-540.

Kellicott, D. S. (1878) A new gall moth and notes on larvae of other gall moths. The Canadian Entomologist 10(11): 202-204.

Miller (1963):
https://kb.osu.edu/bitstream/handle/1811/4921/V63N02_065.pdf

Ming (1989): Thesis, referenced in Foote et al. (1993)

Moffat et al. (2019):
https://link.springer.com/article/10.1007/s10682-018-9966-z

Philips and Smith (1998):
https://www.biodiversitylibrary.org/part/13377#/summary

Phillips (1923): https://www.jstor.org/stable/pdf/25003994.pdf

Steyskal & Foote (1977):
Steyskal, G. C. and Foote, R. H. (1977) Revisionary notes on North American Tephritidae (Diptera), with keys and descriptions of new species. Proceedings of the Entomological Society of Washington 79 (1): 146-155.

Stireman et al. (2010):
https://doi.org/10.1016/j.ympev.2009.09.010

Stoltzfus (1974):
https://lib.dr.iastate.edu/cgi/viewcontent.cgi?article=6120&context=rtd

Stoltzfus (1989):
https://scholarworks.uni.edu/cgi/viewcontent.cgi?article=1587&context=jias

Sutton & Steck (2005):
http://journals.fcla.edu/mundi/article/viewFile/25075/24406

Thompson (1907):
Thompson, M. T. (1907) Three galls made by cyclorrhaphous flies. Psyche 14: 71-74.

Quisenberry (1951):
Quisenberry, B. F. (1951) A Study of the Genus Tephritis Latreille in the Nearctic Region North of Mexico (Diptera: Tephritidae). Journal of the Kansas Entomological Society, Vol. 24, No. 2 (Apr., 1951), pp. 56-72

Posted on October 08, 2019 06:53 by ddennism ddennism | 21 comments | Leave a comment

September 11, 2018

Two Texas Toadshades

Trillium gracile, and T. ludovicianum may both be present in eastern Texas. These are my notes for distinguishing them from one another, taken from the monograph where John Freeman first described T. gracile and clarified the previously-hazy description of T. ludovicianum. Where metrics are given, I have omitted the extreme values, providing only the "typical" ranges (not in parentheses). Other Trillium also occur in Texas. By "leaves" I mean, technically, "bracts".

T. gracile T. ludovicianum
scape
scape length : leaf length 3.2 - 3.5 2.4 - 2.8
leaf
shape elliptic
(elliptic-ovate to elliptic-oblong)
lanceolate or
broadly ovate
sepal
length 18 - 26 mm 24 - 40 mm
apex shape acuminate-blunt or obtuse acute or acute-rounded
petal
length 21 - 35 mm 35 - 55 mm
stamen
anther sac
dehiscence
introrse latrorse
pollen color creamy - yellow olive - orange
anther connective
prolongation
short-beaked short-rounded
gynoecium
carpel-height : stamen-length barely exceeding 0.5 ≥ 0.75
ovary x-section 3-angled 6-angled
stigma length 2.0 - 4.5 mm 4.5 - 10 mm
stigma attachment divergent
extending ovary outline into lyre-shape
initially continuous in outline with ovary
bending distal to point of attachment
fruit
outline smooth often prominently 6-ridged

I have seen comments suggesting that clump-formation and overall plant size differ between the two species, with T. ludovicianum being both more likely to form clumps and generally taller. These are not mentioned in Freeman's monograph. He gives an overlapping range of scape lengths for the two species, with T. gracile being actually taller on average (though it has petals that are about 2 cm shorter):

Trillium gracile: (16-) 20-32.5 (-36) cm long scapes
Trillium ludovicianum: (10-) 15-28 (-37) cm long scapes

Singlehurst et al. (2003) summarized the occurrences of Trillium in Texas. Changes from Freeman's monograph germane to their separation by morphology:

  1. T. gracile is listed as having elliptic to broadly ovate leaves.
  2. T. ludovicianum is listed as having clump-forming tendencies lacking in T. gracile

The two species are probably at least partially temporally isolated with T. gracile blooming later: late Mar -May vs. late Feb - early Apr.

Posted on September 11, 2018 19:03 by ddennism ddennism | 0 comments | Leave a comment

August 10, 2018

Uvularia

U. grandiflora U. perfoliata U. sessilifolia U. puberula U. floridana
leaf attachment perfoliate perfoliate sessile sessile sessile
tepal planar curvature whole tepal twirls along long axis, apex may flare tepal margins may roll outwards, apex may flare apex may flare apex may flare tepal twirls distally along long axis, apex may flare
tepal abaxial surface smooth raised orange-yellow bumps smooth smooth smooth
stem cross-section at nodes terete terete angled angled, with rough puberulence along ridges angled
leaf abaxial surface pubescent glabrous-glaucous glabrous-glaucous puberulent(-glabrous) glabrous

U. floridana also has a leaf-like bract (very, very near the flower) that is absent in its close relatives (U. sessilifolia, U. puberula)

Posted on August 10, 2018 19:25 by ddennism ddennism | 1 comment | Leave a comment

June 26, 2018

Two Erigeron in PA

Erigeron annuus and Erigeron strigosus are the two Erigeron species in PA with tapering leaf bases (rather than clasping leaves).

Flora of PA treatment uses vague descriptors to distinguish them (what constitutes "numerous leaves"?) that might be useful once the user is already familiar with the two.

Flora of North America entry (Guy Nesom, 2004) quickly distinguishes them from other Erigeron based on lack of pappus on ray florets, but not disc florets . This is pictured in my observation.
http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=112000&key_no=5
It then emphasizes stem vestiture, which might not be visible in many observations on iNaturalist. Even this emphasis includes quite a bit of overlap, apparently to accommodate some varieties of E. strigosus and to accommodate the fact that E. annuus sometimes has strigose hairs too.

Michigan Flora gets around this problem by ignoring the problematic E. strigosus var. septentrionalis (associating it with some forms of E. annuus), which makes for a cleaner couplet, but who knows? it might be an oversimplification:

"E. annuus: Middle region of stem glabrate to pubescent with all or many of the hairs long (0.5–1.2 mm) and spreading; principal cauline leaves usually elliptic to ovate, ca. 10–35 (–40) mm wide, with a few large teeth.

E. strigosus: Middle region of stem moderately to densely pubescent with only short (0.5 mm or less) mostly appressed-antrorse hairs; principal cauline leaves linear to oblanceolate, ca. 2.5–10 (–15) mm wide, entire."

https://michiganflora.net/genus.aspx?id=Erigeron

Posted on June 26, 2018 00:34 by ddennism ddennism | 2 observations | 0 comments | Leave a comment

March 28, 2018

Two Blue Cohosh Species

Caulophyllum thalictroides and Caulophyllum giganteum are separate species according, as far as I can tell, mostly to this paper. However, it occurs to me, reading this paper, that physiological effects of early emergence could confound the determination of the morphological details used in this study.

For example, if the emergence and flowering phenology of C. giganteum is earlier, and if the vegetative characters that supposedly distinguish the species continue to expand and grow, as do many forest herbs, then the vegetative characters could appear larger for supposed C. giganteum plants just as a consequence of their head-start. This could be a problem as long as all the plants in this part of the study were sampled on the same day-of-the-year, rather than day-since-emergence (they were).

"On the collection date of the mass sample, 11 May 1982, C. giganteum had completed flowering while C. thalictroides was in anthesis." But how were such plants assigned species-identifications, then? Hopefully not by the same morphological characters that were used in the PCA!

The vegetative morphological characters in the single-population experiment:

Vegetative differences in C. giganteum (all longer and/or bigger):

  1. leaflet length and width of the first two leaves
  2. leaflet sinus length of the first two leaves
  3. primary petiolule length of the first two leaves
  4. terminal inflorescence length
  5. and a decrease in the degree of compounding of the second leaf.
The authors also show that flower size differences distinguish the species. (This part is from herbarium specimens across the ranges of all three species in the genus.)
  1. stamen length
  2. sepal length
  3. pistil length
  4. petal length
  5. ratio of filament length to anther length
    (but you shouldn't use ratios in this type of analysis)

In this case, they found convincing evidence of a bimodal distribution along PCA1 (composed of the above 5 characters, in decreasing order of importance, and with the same positive valence), which suggests two morphologically distinct species, one big-flowered and one small-flowered. However, this shows no evidence of the claimed phenological separation, and doesn't really show evidence of other traits that supposedly differentiate the species (flower number per inflorescence, perianth color). Herbarium specimens are not always the most representative examples of a given population, and there may well have been plants in the C. giganteum populations that had smaller flowers that were less conspicuous to the collectors.

A common greenhouse experiment might be necessary to determine whether there really is separation here, and I'd like to see evidence that organ expansion has completed by the time of its determination in the first part of this study. But maybe first I should observe some of these populations for myself:

The closest Caulophyllum locations to me:
https://www.inaturalist.org/observations/2000688

For my late April trip to Shenandoah:

on the way down:
https://www.inaturalist.org/observations/8032969
https://www.inaturalist.org/observations/3807980
https://www.inaturalist.org/observations/7097471
https://www.inaturalist.org/observations/838935
https://www.inaturalist.org/observations/7646865

in and around Shenandoah:
https://www.inaturalist.org/observations/5100770
https://www.inaturalist.org/observations/3885367
https://www.inaturalist.org/observations/6133924
https://www.inaturalist.org/observations/4799201
https://www.inaturalist.org/observations/5645425

Posted on March 28, 2018 19:43 by ddennism ddennism | 4 comments | Leave a comment

March 16, 2018

Three Similar Toadshades in SE USA

Trillium decipiens, T. reliquum , and T. underwoodii form a group of three closely related species. Here are my notes for distinguishing them from one another, taken from the monograph where the other two were first segregated from a broader concept of T. underwoodii.

T. decipiens T. underwoodii T. reliquum
stalk (scape)
carriage
erect erect decumbent*
stalk : leaf
length ratio
2.5 - 3.0
tall-appearing,
leaf tips don't touch ground
1.0 - 2.5
short-appearing,
leaf tips often touch ground
1.6 - 2.0
short-appearing,
leaf surfaces at or near ground level
sepal
carriage at flowering
divergent-spreading horizontal,
or curving back down to touch leaves
variable
length : width ratio 3.0 - 3.5 3.5 - 4.0 3.5 - 4.0
petal
shape
broadly oblanceolate - obovate narrowly oblanceolate to narrowly elliptic usually narrowly elliptic,
but variable
length : width ratio 2 - 3 3.5 - 5 3.5 - 4
occasionally broader
color highly variable, from
green to yellow-purple to brown-purple
brown-purple
(rarely yellowish)
brown-purple
bronze
(rarely yellowish)
anther sac dehiscence lateral lateral introrse
stamen : carpel
height ratio
≈ 1.5 ≈ 1.5 ≥ 2
leaf shape lanceolate
(straight line from widest point to apex)
lanceolate
(straight line from widest point to apex)
broadly elliptic
(convex curve from widest point to apex)

*not nearly as decumbent as Trillium decumbens. In the T. reliquum population I visited, the stalk (scape) will sometimes only hint at 'laxness' with a slight 'S' bend, but at least some plants in a given population should have scapes that grow initially along the ground.

Posted on March 16, 2018 23:25 by ddennism ddennism | 2 observations | 0 comments | Leave a comment

February 21, 2018

Distinguishing Yellow Trout Lilies in E USA

It's easy to distinguish our three yellow trout lilies from one another when they are in-fruit, or dug-up. But what about when they're in-bloom? You know, when you notice them?

The presence of stolons can be inferred from the number of one-leaved, 'sterile' plants in a population. The stolon-producing species often produce carpets of plants in this stage; E. umbilicatum subsp. umbilicatum will only produce the occasional cluster of steriles, which are either same-aged siblings (clustered by a single fruit dispersal event) or offsets.

Clifford Parks and James Hardin (1963) carried out an exhaustive study of their floral characteristics and correlated them to stolon production, ploidy, and capsule shape. I thought the results of their paper might be useful to iNaturalists. They are summarized here:

E. rostratum E. americanum E. umbilicatum
subsp. umbilicatum
E. umbilicatum
subsp. monostolum
tepal carriage agape strongly reflexed strongly reflexed strongly reflexed
flower angle erect nodding nodding nodding
stolons 1+ 1+ 0 1
capsule shape
in profile
strongly beaked
("rostrate")
rounded, truncate,
or apiculate
indented
("umbilicate")
indented
rarely merely truncate
capsule presentation held erect not erect
but still held off the ground
reclining on the ground reclining on the ground
or rarely just above
petal bases clearly auricled
encircling filaments
minutely auricled
or toothed
not auricled not auricled,
but margin irregular
green coloration
on abaxial side of tepals
none none or slight none present
pale spot
at base of inside of tepals
absent absent in 90%
otherwise vague or small
always present,
but sometimes small
always present
often prominent and large
dark flecking
on perianth
absent absent or slight absent or slight,
but variable
always present,
few to many
style thickness
just below point of
stigmatic divergence
thickened thickened remains thin remains thin
stigma lobes swollen
short
erect
swollen
long
divergent
slender
short
divergent
slender
long
divergent
anther & pollen color yellow
always
yellow
or brown-lavender
brown-lavender
rarely yellow
brown-lavender
rarely yellow
ploidy diploid tetraploid diploid diploid

Not included: E. americanum subsp. harperi, because the authors questioned its distinctiveness. It's mainly distinguished from E. americanum subsp. americanum by having (1) more strongly-apiculate capsules and (2) stigma lobes that are 'distinctly grooved distally' and variously described as 'recurved' or merely 'divergent'. It is documented from Alabama, Georgia, Mississippi, and southern Tennessee. Geraldine Allen and Kenneth Robertson consider it to be more reliably distinct and single it out in their treatment of the genus for The Flora of North America entry.

Posted on February 21, 2018 21:28 by ddennism ddennism | 4 comments | Leave a comment

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