The Amateur Anthecologist

Andrena: A Closer Look

noreply@blogger.com (Travis Owen) — Fri, 18 Mar 2022 11:06:00 +0000

Spring is here! And with it, a new season of bees. Honey bees, having spent the winter clustered for warmth and feeding on honey stored from the previous year, are well adapted to take advantage of the earliest of blooms on sunny days, especially with the disconcertingly dry winter we've had in the Rogue Valley. Bumble bee queens are also out and about looking for potential nest sites, and looking for nectar to replenish them after their long winter dormancy. Despite the widespread recognition by the lay observer of the honey bees and bumble bees, the most speciose group of early spring bees are the mining bees in the genus Andrena.

Andrena males on Polemoneum carneum near Pilot Rock (Ashland, OR)

There are roughly 550 species of Andrena in the United States and Canada, with at least 1500 species worldwide. At least 200 species are found in the Pacific Northwest. Andrena are found all around the world except Australia, South America, and Antarctica (there are no bees in the Antarctic circle). Most species are active in early spring, and there are a relatively small selection of species active in autumn. They are typically small, yet some species are roughly the size of a honey bee. The related species Perdita are the worlds smallest bees with the highest diversity in the Southwestern US and Northwestern Mexico. Some species of Perdita are smaller than mosquitos! The Andrenidae, the taxonomic family that contains Andrena, Perdita, and many other genera is the largest taxonomic bee family with 45 genera worldwide.

Two different sized Andrena females on Ceanothus integerrimus

The majority of Andrena species are univoltine, where there is only one generation per year. Some species are bivoltine with two generations per year. Most species go through their dormant period (winter in North America, though there are winter-active species overseas) in diapause as adults, which likely gives them the advantage of being able to emerge quickly once conditions are favorable. If environmental conditions are not ideal for the flowering plants they depend upon, some species have been known to remain in diapause for at least two years before emerging.

Andrena pertristis (?) on common dandelion

Most Andrena are solitary, every female excavates and provisions her own nests. Some species are communal and dozens of females will share a single nest entrance, but they still lay eggs in their own cells inside the communal nest. It's comparable to a New York apartment building, where neighbors share a front entrance but still have to buy all their own groceries. Some Andrena nest in large congregations where nest entrances may be as close as an inch apart. Nest entrances may be hidden under leaves or fallen branches, and nests may be excavated in various soil types (i.e. sand or clay).

Male Andrena perched on Fritillaria affinis, near Pilot Rock

Unlike honey bees, Andrena excavate tunnels straight down into soil. They are very common in lawns and sometimes nest in soil covered by vegetation, unlike most other ground dwelling bees. Cells branch off the main shaft on short lateral tunnels. Nests of Andrena are typically between 6-12" (15-30cm) deep, but desert dwelling species nests can be incredibly deep. Andrena, like many solitary ground nesting bees, line their cells with a wax like substance produced from the Dufour's gland on the underside of the abdomen, and spread it around the cell wall with the trowel-shaped pygidial plate located at the tip of the abdomen. The waxy substance protects the larvae from bacterial infections and retains the moisture in the cell, reducing the risk of desiccation. Andrena also use the Dufour's gland secretions to mark the entrance of the nest to help them find the nest via the odor, especially important in communal nests where keeping intruders out is of great concern. Additionally, female Andrena scent-mark flowers with Dufour's gland secretions where resources were good so they may return on the next foraging trip.

Andrena on February blooming Scilla mischtschenkoana in my garden

Early spring weather is sometimes unpredictable. Rainy and windy weather can make it difficult for early bees to forage for resources and provision cells. With all solitary bees, cell provisioning entails collecting pollen and nectar, mixing them to form a pollen ball, depositing an egg, and sealing the cell before starting on the next one. Andrena are typically slow to provision cells, sometimes only completing a single cell per day. On average, each foraging trip can take over an hour for some species to gather nectar and pollen for a cell, and an average Andrena cell takes roughly nine foraging trips to complete. Some of these foraging trips are used for the mother bee to feed on nectar for herself. Nectar and to a lesser extent pollen is the sole source of carbohydrates and nutrients for adult bees. Pollen is the sole source of proteins and amino acids bees need, especially important for developing larvae.

Lomatium californicum on Upper Table Rock with swarming male Andrena

Andrena are short tongued bees, and most forage for nectar on flowers with easily accessible nectar. Unlike honey bees and bumble bees which use their proboscis more like a straw, short tongued bees lap up nectar not unlike a cat. This doesn't mean all Andrena can't access nectar in larger or deeper flowers, some species have longer mouth parts or special adaptations that enable them to reach hidden nectar. Many species are pollen specialists and only collect pollen from a narrow range of plant species. For example, some Andrena species only collect pollen from Asteraceae (sunflower family), Apiaceae (parsley family), or willows (Salix spp.). The specialists will still collect nectar from a wider range of plants, including many nonnative plants. Most species aren't terribly picky, and some are important pollinators of cultivated plants in gardens and agricultural settings, though mining bees aren't commercially managed like alfalfa leafcutter bees or orchard mason bees.

Male Andrena with extended proboscis

Like many of the solitary bees, male Andrena emerge from diapause before the females. Males do not actively collect pollen, but they do feed on nectar. While male bees are not usually often valued for their pollination services, male Andrena do serve important roles in the pollination of some plants like bee-mimic orchids (Ophrys spp.) in Eurasia and Northern Africa where the orchids live. The bee orchids are only pollinated by male bees who are duped into thinking the orchid is a female ready to mate, and they recieve nothing for their efforts. For most other plants, males may be superior pollinators on a per bee basis. Female Andrena collect much larger amounts of pollen than males, but most of that pollen goes to the nest instead of the next flower since females frequently and thoroughly groom the pollen into their scopae (specialized pollen carrying hairs). Males don't groom frequently, and on a per bee basis likely deposit more pollen than females. Males don't visit nearly as many flowers as females, and are generally smaller in size. The importance of any pollinator to a specific plant species depends on how effective each flower visit is in regards to collecting or depositing pollen, and the rate of visitation. On a net basis, females are more important pollinators for the majority of plants visited.

Plum blossoms with an Andrena male

Females collect pollen by grabbing onto the anthers with their legs and bending the abdomen to push the anthers onto their bodies. On some flowers they contact the anthers by default as a result of reaching for nectar. Female Andrena are usually active for much of the day when weather is favorable, and visit many more flowers than males. Between flower visits, female Andrena groom all of the pollen onto their hind legs. Scopae, the specialized plumose hairs used for holding pollen, are located on the hind leg tibia and extending all the way up to the trochanter and propodeum (where the hind leg meets the thorax). Andrena with a full load of pollen can appear to have a belt of pollen spanning between the hind legs.

Female Andrena with pollen laden scopae

Flower visits by Andrena aren't always mutually beneficial. Some Andrena are nectar thieves of certain flower types, accessing the nectar without contacting the anthers or stigma and thus being of no benefit to the plant. Nectar is a metabolically expensive incentive plants make that is meant to attract pollinators to facilitate plant reproduction. Some plants keep the nectar hidden within parts of the flower inaccessible to some pollinators. This excludes the least effective pollinators for a particular plant species and encourages the best pollinators for the particular flower type (or at least that's how it's supposed to work). Many bees cheat by making incisions in the sides of the flowers to bypass the reproductive organs altogether and access the nectar directly.

Male nectar thief stealing nectar from a slit in the side of a manzanita flower.

In my observations, male Andrena appear to steal nectar more often than females. With shallow flowers that don't exclude pollinators, like many of the Asteraceae and most fruit trees, this doesn't appear to be the case. Flowers on plants in the Ericaceae (i.e. Arctostaphylos; Arbutus), Berberidaceae (Epimedium), or Fumariaceae (Corydalis; Aquilegia), to name a few examples, often have either restricted openings in urn-shaped flowers or nectar filled spurs are often victims of nectar thievery. In the case of Arctostaphylos, and many other genera in the Ericaceae, the flowers are only pollinated by bees capable of sonication where the bees vibrate their wing muscles to release the pollen.

Manzanita flowers

Manzanitas, the common name for Arctostaphylos, are one of the earliest blooming shrubs in the Rogue Valley, and they attract a wide range of pollinators, including many hungry Andrena. Bumble bee queens and anthophorine bees (Eucera sp.) use their mandibles to cut slits at the base of manzanita flowers. Smaller bees like Andrena then follow in their place. Most Andrena are not equipped to access the nectar from the flower opening unless the flower is damaged or malformed, and it is unlikely they contribute towards the pollination of Arctostaphylos in any significant way.

Lasthenia californica ssp. californica and male Andrena on Upper Table Rock

Despite being occasional nectar thieves, Andrena are very important early pollinators for many plant species. In some areas, they are the most abundant bee when the weather is cool. Although they are active in early spring in low elevations, they are active in early summer in higher altitudes such as subalpine and alpine mountainous environments. In either high or low elevations, Andrena can't warm their bodies the way that bumble bees do by vibrating their flight muscles and burning calories for warmth. All bees are ectothermic (cold blooded), while some bees, like bumble bees, are facultatively endothermic (having the ability to warm themselves). To cope with cool environmental conditions, Andrena sunbathe on sunny vegetation until their body temperature is adequate for flight.

Fritillaria affinis with possible Andrena pollinator?

Most Andrena feed on nectar and pollen from a very wide range of plant types. While there are many specialist species, most are highly adaptable generalists. Species whose ranges span multiple ecoregions (such as A. prunorum which is found throughout most of the west in the United States and Canada) are not picky eaters, and may feed on completely different plant species in different parts of their range. There are species that collect pollen from many wind pollinated plants, like grasses and oak catkins. Planting flowers for Andrena in the garden is not challenging, so long as you include many species with open or small flowers. Some of the plants in my garden that I have observed Andrena visiting include bulbs (i.e. Crocus; Scilla), strawberries, Apiaceae (Lomarium; others), California lilac (Ceanothus), Berberidaceae (Epimedium; Mahonia), and many others. It's worth experimenting and observing to see what they like, but if you include a large diversity of flower shapes and overlapping bloom periods, there are good odds you'll see some Andrena.

Andrena female gathering pollen on oak catkins, I was very excited to capture this photo!

If you don't detect any Andrena in your area, they may be detected by the presence of some of their parasites. The fuzzy bee-like beefly (Bombylius) is a nest parasite of many ground nesting bees. They have a long needle-like proboscis and visit many of the same flowers as Andrena. Beeflies hover over the ground looking for bee nests, and when one is detected, they skillfully toss some eggs into the nest with a flick of their abdomen, all while remaining in flight. Another group of nest parasites are the cuckoo bees in the genus Nomada. Nomad cuckoo bees are very wasp-like in appearance, often in bright colors like red. While these waspish bees don't collect pollen, they feed on many of the same nectar plants as Andrena and may play a small role in pollination. There are others, and the presence of any of these parasites is a sign of a healthy host population.

Nomad cuckoo bee (Nomada)

I hope you enjoyed reading this introduction to the mining bees, Andrena. This wasn't meant as an identification guide, and there is much more that I could have written to further illuminate the biology and behavior of this large genus. If you would like to learn more, please take a look at some of the resources below.

Sources:

Danforth, Bryan. The Solitary Bees: Biology, Evolution, Conservation. 1st ed., Princeton University Press, 2019.

Fowler, J. (2020). Pollen Specialist Bees of the Western United States. https://jarrodfowler.com/pollen_specialist.html

Michener, Charles. The Bees of the World. 2nd ed., Johns Hopkins University Press, 2007.

Miliczky, Eugene. “Observations on the Nesting Biology of Andrena (Plastandrena) Prunorum Cockerell in Washington State (Hymenoptera: Andrenidae).” Journal of the Kansas Entomological Society, vol. 81, no. 2, 2008, pp. 110–21.

O’Toole, Christopher. Bees: A Natural History. First Edition, Firefly Books, 2013.

Stephen, William Procuronoff, et al. The Biology and External Morphology of Bees. Amsterdam University Press, 1969.

Tang, Ju, et al. “Pollinator Effectiveness and Importance between Female and Male Mining Bee (Andrena).” Biology Letters, vol. 15, no. 10, 2019, p. 20190479.

Wilson, Joseph, and Olivia Messinger Carril. The Bees in Your Backyard: A Guide to North America’s Bees. Princeton University Press, 2015.

Bumble Bees of Vesper Meadow

noreply@blogger.com (Travis Owen) — Thu, 27 May 2021 20:39:00 +0000

Bombus vandykei on a native Phacelia

Bumble bees are some of the most conspicuous of the bees. Fast flying, buzzing balls of fuzz hurtle from flower to flower and can bee seen, or heard, from a distance. This may be part of why they are many folks favorite bees. While I could never choose a favorite bee, bumble bees are charismatic and fascinating. While the United States boasts around 40 species of Bombus (the genus to which bumble bees belong), there are roughly 30 species native to just the Pacific Northwest (Idaho, Oregon, and Washington). At least 15 species call Southern Oregon home, and many of these have been identified from the Vesper Meadow Restoration Preserve in Ashland.

Prime bumble bee habitat: Wyethia angustifolia blooming in part of the Vesper Meadow Restoration Preserve

Vesper Meadow Restoration Preserve

Vesper Meadow is a privately owned 403 acre upland wet meadow in the heart of the Cascade-Siskiyou National Monument. The Vesper Meadow Education Program, which operates at the meadow and surrounding forest acreage, brings together scientists, students of all ages, and the public to study the local ecosystem and cultural significance of the land. One of the goals of the Preserve is to monitor wildlife of all sorts, from birds to plants to insects. Pollinators, especially butterflies and bees, are also being documented and monitored by organizing focused bioblitzes. Typically taking place over the course of a single day, a bioblitz is when a group of volunteer citizen scientists attempt to record as many living species as possible in a given area.

Bombus vosnesenskii on Solidago at Vesper Meadow
2021 Bumble Blitz at Vesper Meadow

Vesper Meadow is hosting a targeted bumble bee survey on Sunday, June 27th (from 10:00am to 2:00pm). This will be the second annual Bumble Blitz. Attendees will learn the basics of bumble bee identification, how to net a bumble bee, become acquainted with some of our native flora, and contribute to science! Also, it's beautiful there. Get some fresh air, meet fellow bee nerds, and enjoy the wilderness. If you plan on attending, please RSVP on the Vesper Meadow website.

Bombus flavifrons on Agastache urticifolia

Bumble Bees 101

I will now provide a basic introduction to some of the bumble bees we've identified from Vesper Meadow. But first, a little about bumble bees. Bumble bees (genus Bombus) are cousins of the honeybees (genus Apis). Bumble bees are found around the world in temperate regions, most north of the equator (with the exception of South America where a handful of species are found), with the highest species richness from the Tibetan Plateau westward to the Alps. Most species are eusocial, like honeybees, except a smaller proportion of species which are social parasites (subgenus Psithyrus, they expropriate established nests of non-parasitic species for their own reproductive gains). In contrast, honeybees aren't native to the Americas and the genus Apis contains no social parasites.

Bombus californicus on Wyethia angustifolia
Bumble Bee Life Cycle

Bumble bee colonies have an annual lifecycle. At the start of the season (anywhere between spring and summer, exactly when depends on the species) a queen emerges from her hibernaculum and seeks both nourishment from flowers and an abandoned rodent burrow or grass tussock to initiate a nest in. Once she finds a suitable nest location, she rears a batch of miniature workers which take over some of the foraging duties of the queen, allowing her to remain in the nest more and more. Eventually the colony has many workers (between 50 and 500, though usually less than 100) and the queen remains in the nest for the rest of her life. Towards the end of the lifecycle of the colony, new queens and drones (males) are produced, which leave the nest to find mates from other colonies. Following mating, the drones die and the newly mated queens seek a place to shelter over winter. The original colony and queen dies, and only the new queens last the following year to renew the cycle.

Bombus bifarius s.l. (B. vancouverensis nearcticus)
Vesper Meadow Bumble Bees

I've caught at least seven species of Bombus at Vesper Meadow, in addition to perhaps three or four additional species which have yet to be identified. Bombus californicus (syn. fervidus), flavifrons, melanopygus (syn. edwardsii), mixtus, nevadensis, vancouverensis nearcticus (syn. bifarius s.l.), and vosnesenskii have all been identified, yet caliginosus and vandykei (very close mimics of vosnesenskii) may possibly be found at or around Vesper Meadow. The white-shouldered bumble bee, B. appositus, is also found in the area (likely at Vesper Meadow) but it is uncommon. Additionally, there has been a single sighting of the Western bumble bee, B. occidentalis, but no sightings since to my knowledge.

Bombus flavifrons

There also may be one or two of the socially parasitic bumble bees, subgenus Psithyrus, though no sightings or specimens have been confirmed as of this writing. The Oregon Bee Atlas expects to find the Fernald cuckoo bumble bee (B. fernaldae, syn. flavidus) and the Indiscriminate cuckoo bumble bee (B. insularis) in Oregon, but very few specimens have been vouchered. Social parasitic bumble bees have only two castes, female ("queens") and male, unlike other bumble bees which have queens, workers, and males. Rather than building their own nests, Psithyrus females (not technically queens since there is no worker caste) invade established nests and take them over, using the host workers to rear her young. This results in killing the colony since workers are no longer produced, only reproductive males and females of the social parasites. This seems harsh, but their presence would indicate a healthy host population. Psithyrus females can be identified by having no corbiculae.

Species Accounts

The following are a selection of bees I've photographed or collected for the Oregon Bee Atlas and Vesper Meadow Bumble Blitz. For the Bumble Blitz, we will be using the capture, chill, release method. This involves netting bees, putting them in vials, and chilling them in a cooler so we can take detailed photos of them for identification. This is in contrast to netting bees, placing them in a kill jar (it's more humane than it sounds), and pinning them for identification under a microscope. Both collection methods have pros and cons. There are some bees that are very difficult to identify in the field, which is where pinned specimens may be superior. However, catch-and-release doesn't come with any moral ambiguity. As you will notice, photos don't always capture every characteristic necessary to assuredly identify some species. The best advice is to take as many good photos as possible, and hope you've captured the right angles.

White-Shouldered Bumble Bee - Bombus appositus

Bombus appositus

The white-shouldered bumble bee, Bombus appositus, has a general appearance of being well groomed with short, even hair all over. As the common name suggests, this bee has white patches on the anterior of the thorax, with a black band between the wing bases. The face and posterior of the thorax is either white or yellow, with the abdomen completely yellow or tawny buff. The white-shouldered bumble bee nests both underground and on the surface, such as within dense grass tussocks.

California Bumble Bee - Bombus californicus

Bombus californicus

Bombus californicus (syn. B. fervidus californicus) is very similar to both vosnesenskii and vandykei, all are black with yellow bands on the thorax and a yellow band of hair towards the tip of the abdomen on the third or fourth segments (also called tergites). B. californicus differs from the lookalikes by having a completely black head and a cheek longer than broad. They nest on the surface or aboveground, occasionally underground. Nests may be found in haystacks, abandoned mouse nests, or birdhouses. They are said to be one of the more aggressive species, most likely when nests are disturbed.

Yellow-Fronted Bumble Bee - Bombus flavifrons

Bombus flavifrons female

Bombus flavifrons females are usually characterized by yellow and black hairs intermixed on the face, and intermixed yellow and black on the anterior of the thorax. The first two tergite have yellow bands while the third and forth tergite have either orange or black. The forms I've seen at Vesper Meadow usually have black bands with few orange hairs intermixed, appearing dull orange. Similar to californicus, they have a relatively long cheek (malar space longer than wide).

Bombus flavifrons male

In contrast to females, male flavifrons are almost entirely yellow or sometimes have black or orange hairs intermixed in the last few tergites. This species nests primarily underground, utilizing abandoned rodent burrows.

Fuzzy-Horned Bumble Bee - Bombus mixtus

Bombus mixtus

Bombus mixtus has yellow and black hairs intermixed on the face and thorax. The first abdominal tergite is always yellow, followed by a black band and the fourth/fifth tergites orange, sometimes appearing dull and intermixed with yellow hairs. B. mixtus is sometimes known as the fuzzy-horned bumble bee because the males have tufts of hair jutting from their flagellum (segments of the antennae), though they're only visible at certain angles under magnification. They are flexible in their nesting locations, nesting above- and belowground.

Black-Tailed Bumble Bee - Bombus melanopygus (syn. edwardsii)

Bombus melanopygus (syn. edwardsii)

As seems to be the case with many of the Western bumble bees, Bombus melanopygus has two distinct color morphs. The morph pictured here has no orange, and is sometimes considered a separate species, B. edwardsii, depending on the authority. The black-tailed bumble bee, B. melanopygus, is so named since the last tergite is always black. The hairs on the head are usually yellow or yellow intermixed with black. Similarly, the anterior of the thorax is mostly yellow with black intermixed, followed by a black band, with yellow hairs on the thorax posterior. Note that the corbicular fringes are mostly black (compared to vancouverensis nearcticus which has orange corbicular fringes) The abdomen always has yellow on the first tergite, usually followed by orange on the second and third tergites.

Bombus melanopygus (syn. edwardsii)

The Bombus edwardsii color form of melanopygus replaces orange with black on tergites two and three. The fourth and fifth tergites have yellow creeping inwards from the sides. B. vancouverensis nearcticus is very similar to the edwardsii color form, but nearcticus (almost) always has black on the last two tergites in females. The black-tailed bumble bee nests wherever, in birdhouses, housing insulation, abandoned rodent burrows, or wherever they deem fit.

Nevada Bumble Bee - Bombus nevadensis

Bombus nevadensis queen

Bombus nevadensis is similar to B. griseocollis which is more common at lower elevations. B. nevadensis has a look of being clean cut. It seems more brown than yellow compared to other bumble bees in the area. Hair on the face is usually black, sometimes with a few yellow hairs intermixed. There is usually a black spot or band between the wings with many intermixed yellow and black hairs around it. The sides of the thorax are black. The second and third tergites always yellow, while the first tergite is either black or yellow. The last few segments are always black, except for males which have orange at the tip. These large bees are most often ground nesters.

Two-Formed Bumble Bee - Bombus bifarius s.l. (syn. vancouverensis nearcticus)

Bombus vancouverensis nearcticus (syn. B. bifarius s.l.)

Bombus vancouverensis nearcticus (sometimes shortened to B. nearcticus) and v. vancouverensis were recently split from bifarius, the two-form bumble bee (see Ghisbain et al 2020). According to genetic analysis, bifarius s.s. is restricted to the Colorado Plateau and surrounding regions east of the Rockies, while vancouverensis nearcticus is found from the San Juaquin Valley in California and as far north as British Colombia. The third species, v. vancouverensis is restricted to Northwestern Washington and Southwestern B.C. If you don't follow, don't worry. Oregon only has v. nearcticus, which has no orange on the abdomen, while both bifarius s.s. and v. vancouverensis both have red.

Bombus vancouverensis nearcticus

Bombus vancouverensis nearcticus has a characteristic black notch in the center of the yellow band on the posterior of the thorax. There is a yellow band on the fourth tergite while the fifth and sixth tergites are always black. Compare with edwardsii (syn. melanopygus) which has yellow on the sides of the fifth tergite. This species will also have orange fringes on the corbicula while edwardsii will usually have black hairs, sometimes tipped with orange. The two-form bumble bee is primarily an underground nester, though they occasionally nest on the surface.

Yellow-Faced Bumble Bee - Bombus vosnesenskii

(including descriptions for vandykei and caliginosus)

Bombus vosnesenskii

Bombus vosnesenskii, vandykei, and caliginosus are all incredibly similar. All three have yellow faces and yellow shoulders, with a yellow band on the abdomen. With vosnesenskii and caliginosus, the yellow abdominal band will be on the fourth tergite while vandykei will have the yellow band on the third tergite. Where in vosnesenskii the sternum (underside of the abdomen) will be entirely black, caliginosus will have pale yellow fringes on the fourth sternite. I've also noticed that vandykei males are often completely yellow and probably indistinguishable from male flavifrons. Both vosnesenskii and vandykei nest underground, while caliginosus will nest underground or in abandoned bird nests.

- - -

Thanks for taking the time to read this. This is no substitute for a proper identification guide, but rather an attempt at an introduction. I hope you can make it to the 2021 Bumble Blitz, so if you can please RSVP!

Learn more about Bumble Bees:

The Bumble Bees of the Oregon Bee Atlas - Lincoln Best (YouTube)

PNW Bumble Bee Atlas - How to Identify Pacific Northwest Bumble Bees - Xerxes (YouTube)

PNW Bumble Bee Atlas (Website)

Genus Bombus - BugGuide.net (Website)

Guide to Bumble Bees of the Western United States - USDA (PDF)

Table Rock Revisited

noreply@blogger.com (Travis Owen) — Thu, 08 Apr 2021 06:44:01 +0000

Spring is the best time of year to hike up either of Southern Oregon's Table Rocks. The sky is blue and fresh, life has returned to the land after a grey winter, and the humming of insects fills the air. Flowers are the jewels of the experience, since they are the epitome of Spring.

A mining bee (Andrena)

Part of what makes Upper Table Rock enjoyable to hike is that it's relatively quick and easy, not to mention very close to where I live. It takes less than an hour to reach the top of the mesa. Of course, with my camera I tend to stop and snap a few photos every time I see something interesting so it often means I spend most of the day exploring.

Erythronium hendersonii

Shortly after setting out on the trail, Erythronium hendersonii takes center stage. It was in bloom as far as I could see in the forest understory on the first half of the ascent. I advise getting on your hands and knees to check out the intricate floral markings! These nectar guides may help bees (the primary pollinators of Erythronium) find the nectar reward at the intersection of the petals.

A carpet of Erythronium

Inexplicably, bees were largely absent from most of the flowers lower on the trail. A few factors may have contributed to this. The manzanita was blooming further up the plateau, and the bumblebee queens which I suspect are the primary pollinators of erythroniums here were busy looking for suitable nest sites. One can only imagine.

Apis mellifera

As I continued to ascend the trail, honey bees became more prevalent on the erythroniums. Perhaps they've replaced the ancestral bumblebee pollinators. Maybe the bees which once pollinated E. hendersonii are now extirpated. Ranges of species change all the time, at least in evolutionary timescales. Many species of bumblebees have been seeing declines in recent decades due to habitat loss and diseases spread by introduced species. This is purely conjecture.

Honey bees appear to be adequate pollinators of E. hendersonii. The stigma is situated in a way that simply jostling the flowers (such as a bee landing on the stamens) may cause some pollen to move around. Most of the erythroniums that I've encountered in the Rogue Valley set seed every year despite potentially low numbers of pollinator visits. This apparent ability to set seed easily may be an adaptation to unpredictable spring weather. It's around this time of year that rain becomes an oddity and the next time water falls from the sky is autumn. Time to dry out.

Protosmia rubifloris ♂ hiding in Ranunculus occidentalis

The abundance and variety of bees now is astonishing. Protosmia is an osmiine bee (Megachilidae, leafcutter, mason, and resin bees) that nests in above-ground cavities. Records of nest sites is sparse, but some species have been known to make nests in things such as pine cones and snail shells! Another trait that makes Protosmia fascinating is that they overwinter as adults, uncommon in the Megachilidae. Their nests are capped with an excreted resin. Another peculiarity is that not all bees favor buttercups (Ranunculus spp.), and the pollen is even toxic to some bees.

Anthaxia aff. inornata

It is likely beetles are often overlooked as pollinators. They're not as efficient, or perhaps charismatic, as bees. Beetles were pollinating gymnosperms like cycads at least 200 million years ago, at least 75 million years before angiosperms (flowering plants, plants most adapted to bee pollination) and bees every appeared in the fossil record. Many flower-visiting beetles don't travel from flower to flower as frequently as bees, but there are some with densely haired bodies which are important pollinators of some plant groups (including many of the large-flowered tulip species), many bearing bowl shaped magnolia-like blossoms.

Flower longhorn beetle (Cortodera)

Beetles visit flowers for a variety of reasons, though I don't know of any beetles that collect pollen for rearing young. Many beetles eat pollen, nectar, or the flowers themselves. In other instances, beetles congregate at flowers to mate. There are also parasitoid beetles which lay eggs on or near flowers, the larvae attaching themselves to bees to hitch a ride to the nest where they feed on the provisions supplied by the bee or the bee larvae.

Ranunculus austro-oreganus

Southern Oregon buttercup (R. austro-oreganus) are endemic to, unsurprisingly, Southern Oregon (specifically Jackson County, according to the NRCS). They are slightly larger than R. occidentalis, and they have reddish veins on the back of the petals. Upper Table Rock hosts a vigorous population on the first half of the trail.

A hidden Lasioglossum ♀ forages on Southern Oregon buttercup.

I'm always intrigued by bees that visit buttercups. Many of the bees that collect ranunculus pollen are specialists, and the pollen would be toxic to many bees unable to metabolize it (Praz et al. 2008). That said, seeing a bee on a buttercup seems like something special. Pollen specialists need the pollen from their host plant in order to rear the next generation, which elucidates why maintaining a healthy balanced ecosystem is so crucial. Lives literally depend on diversity.

Fritillaria recurva

One plant that always results in a gasp is the beautifully checkered Fritillaria recurva, sometimes called red bells. Upper Table Rock sports a scattered population of the gloriously red bulbs. I have never once seen pollinators of any kind visit the flowers, but it isn't terribly surprising since I have never seen more than a dozen of them flowering in one place. Pollinators, particularly bees, are generally most attracted to large patches of flowers of the same species. The most F. recurva flowers I have seen in one place is maybe a little over a dozen. Besides all that, the morphological characteristics of the flowers suggest Ornithophily (bird pollination). Red tubular or bell shaped flowers with dilute nectar and slightly protruding stamens fits the bill. [Pun was accidental]

Fritillaria recurva is one of the most graceful native wildflowers, and it's always a treat to find in the wild. I hope to someday see a hummingbird visiting the flowers, but due to a lack of photos of pollination events on the google, I may be in for a bit of a wait.

Habropoda depressa ♂ on Cynoglossum grande

Another woodland favorite is the Pacific hounds tongue (Cynoglossum grande); a bigger, sexier version of the familiar forget-me-not (Myosotis) grown in cottage gardens. Being a member of the Boraginaceae (you guessed it, the borage family), it has white fornices (the white appendages at the "eye" of the flower, sing. fornix) and hidden anthers. Bees collect pollen by sticking their tongues in the flower, and some have specialized hooked hairs on various parts of the proboscis for this purpose.

Habropoda depressa ♂

Habropoda depressa, sometimes known as mountain digger bees, are some of my favorites. As the name suggests, they are solitary ground-nesting bees. Despite being solitary, they nest in large congregations. Males are the first to emerge, so the first to see foraging for nectar as they wait for the females. As the unfortunate females emerge a week or two after the males, they are balled by thirsty males trying to mate. I observed such an occurrence recently. The males numbered probably a dozen per female, and females couldn't even get off the ground before being mobbed. This doesn't appear to hurt the bees, despite some grappling between males, and they seem to have very healthy populations here.

Andrena on Lomatium urticulatum

Mining bees (Andrena spp.) are the most common spring bees on Upper Table Rock. Spring gold (Lomatium urticulatum) is the first of the buscuitroots to bloom, sometimes starting in mid February. Small bees and flies are quite smitten.

Formica

Ants are one of the least likely pollinators, despite many species seeking nectar on a huge variety of plants. Their ability to be good pollinators is sometimes hindered by pollen-destroying excretions on the exoskeletons of some species. Many ants produce antibiotic compounds needed to keep pathogenic microorganisms out of the nest. Pollen that comes into contact with these compounds is often no longer viable, and ant pollinated plants see a decreased seed set when compared to bee pollinated plants. See Beattie et al. 1984.

Andrena males on Lomatium californicum

Larger lomatiums grow in clearings along the trail on Upper Table Rock. The inconspicuous green flowers are highly sought by small mining bees. Andrena are short tongued bees, having relatively short labial palpi and glossa (for detailed diagrams of bee mouthparts see Bee Anatomy, it's a European bee ID website but the diagrams are good for all bees). The glossa can be thought of as the true tongue, the part that laps up nectar. In general, short tongued bees need flowers with easily accessible nectar and shallow flowers. Lomatiums and many genera in the Apiaceae (carrot family) offer numerous small flowers which are attractive to many small beneficial insects.

Andrena ♂

Andrena are mostly solitary, but many are thought to be communal. This means the females share a nest entrance but dig and provision their own cells which radiate laterally off the main burrow. Andrena line their cells with a wax-like substance, possibly to keep water or pathogens out. Males are sometimes seen before females, but they both usually emerge at the same time. Both male and female overwinter as adults, though the females mysteriously hide after mating, perhaps returning to the burrow to start digging cells. Andrena are univoltine, meaning one generation is produced per year. More on Andrena later.

Micranthes integrifolia with a fly

Micranthes, formerly Saxifraga, is seen periodically on both Table Rocks. It appears to be attractive primarily to various flies, as I suspect most saxifrages are.

Buckbrush (Ceanothus cuneatus)

At the time of my hike on Upper Table Rock, the buckbrush was just about to bloom. (By thy time I finish writing this, the bloom will probably be passed.) The very first of the musky smelling ceanothus blooms were attractive to honey bees. The only open buds were on the sunny south side of the shrubs. Sunlight is a key indicator to many plants that it is time to bloom, be it day length or plain old warmth.

Birch leaf mountain mahogany (Cercocarpus betuloides)

I've probably walked by these trees in bloom a hundred times in complete ignorance. The greenish inconspicuous flowers of mountain mahogany were attractive to bumble bees and honey bees, and the buzzing was the only indicator to me that they were blooming.

A honey bee grooms pollen into her corbicula

The ten to twenty foot tall trees periodically grew along the Upper Table Rock trail. Mountain mahogany is common in the Western states, growing in dry mountainous habitats from Canada south to Mexico. The seeds are very distinctive, and I've seen them for years not knowing what made them. Mountain mahogany seeds float down with a spiraled fuzzy tail, similar in function to the wings on maple seeds, aiding the seeds in spiraling away from the parent plant. Upper Table Rock trail in the fall is teeming with the peculiar curly-tailed seeds.

California sandwort (Minuarta californica)

Upon reaching the top of the mesa, I was rewarded with a carpet of miniature blooms spanning as far as I could see.

Apis mellifera

Bees were everywhere. I had seed the occasional honey bee, but the vast majority of the bees on the summit were small native bees. Every time this honey bee landed on a flower it bent it over.

Andrena

The majority of bees on the top of Upper Table Rock were small natives, mostly Andrena. These small bees, roughly half the size of a honey bee, are completely silent in flight. Honey bees, in contrast, are noisy and buzz every time they fly. This is a common phenomenon when comparing colony-forming Hymenoptera with solitary Hymenoptera. Colony forming bees (i.e. honey bees, bumble bees) and wasps (i.e. yellowjackets) are noisy and conspicuous, often brightly colored. This is an aposematic warning to would-be predators that they can and will defend themselves.

Andrena

Solitary bees (i.e. mining bees) and wasps (i.e. braconids) are often quiet in flight, and often have dull colors. These bees would rather flee than fight. Compared to honey bees, many solitary bees have relatively little invested in their nests and can readily relocate, abandoning a compromised nest. Nests of honey bees, bumble bees, and yellowjackets (among others) are full of hard earned resources (i.e. wax/paper comb) and multitudes of developing larvae. If these social Hymenoptera were forced to abandon their nests, it may be a death sentence. Solitary bees, like Andrena, aren't well equipped to defend the nest from large predators, so they don't advertise it.

Andrena

Andrena collect pollen dry, without mixing it with nectar like honey and bumble bees do. Specialized hairs called scopae are found on the back legs of Andrena all the way from the tibia all the way up to the propodeum (the rear, bottom portion of the thorax). Pollen appears to wrap around the underside of their waist, like a belt. While this dry collected pollen has a better chance of becoming dislodged on a flower (unlike wet honey bee pollen), it is estimated that less than 5% of the pollen is used for pollination. That means that at least 95% of the pollen collected by bees is eaten. This makes me wonder, do plants even want bees to collect their pollen?

Andrena ♂

Male Andrena were common at the time of my hike. Their long antennae are used to detect potential mates. Males don't collect pollen, and their utility as pollinators is questionable. However, we can thank the males for creating the next generation of Andrena! Thanks guys. Your bee-havior has not gone unnoticed!

Lasioglossum (subg. Lasioglossum)

Sometimes known as the sweat bees, Lasioglossum are probably the most abundant and diverse bees in North America, often very small and inconspicuously colored. They range in sociality from true solitary to primitive eusociality, sometimes varying geographically possibly in response to resource availability. Nearly all Lasioglossum nest in the ground, and some can have multiple generations a year with some overwintering as adults and some overwintering as larvae. There is a lot of mimicry between species which can make identifying Lasioglossum to species very challenging, something I hope to get better at this year.

Lasthenia californica ssp. californica with male Andrena

More Andrena on dwarf daisies on Upper Table Rock, shocking! The common name for Lasthenia, goldfields, is extremely accurate. Large swaths of golden honey-scented daisies carpeted the mesa summit intermixed with California sandwort. Goldfields vary in height from less than two inches up to fifteen inches depending on the soil. The soils atop the Table Rocks is thin and probably deficient in nutrients, resulting in dwarf versions of otherwise foot-tall flowers. L. californica is native throughout the West Coast from Southern California to Southwestern Oregon, mostly on the western halves of the states.

Andrena

It would be interesting to learn whether different species of Andrena prefer different plant species blooming concurrently. I didn't collect any specimens during my hike, but it would be interesting to investigate in the future. Many Andrena are pollen specialists of Asteraceae and even single genera or species. Pollen specialist means the females collect pollen from a narrow range of genera or species, but may collect nectar from a wider array of plants. Goldfields (Lasthenia) are host plants to at least a few species of Andrena (see Thorpe and Leong 1998).

Plagiobothrys aff. nothofulvus

Popcorn flowers, Plagiobothrys spp., are host plants for a ton of native pollen specialist bees in Oregon. I saw many patches of popcorn flower in various parts of the Upper Table Rock hike. I didn't see any bees, but I didn't observe for long. Pollen specialists of many members of the Boraginaceae (the forget-me-not family) have special adaptations for collecting the pollen hidden in the flowers. Some of these bees have velcro-like tongues with specialized hooked hairs on the proboscis. So cool!

Allium parvum

One of the treats I always look forward to on Upper Table Rock are the diminutive onion flowers which sit on the ground, often against volcanic stones. Allium parvum is a tiny onion with pea-sized bulbs. The leaves are often dead or dying by the time the flowers open.

Collinsia sparsiflora

Another dwarf version of an otherwise taller species, the odd-named spinster's blue-eyed Mary. In better soils, these plants can grow up to a foot tall, though on the thin rocky soils of Upper Table Rock they are less than two inches. Small three-square foot patches variously carpeted the mesa summit. Oddly, I witnessed no bees on these flowers. Peculiar.

Cowbag clover (Trifolium depauperatum var. depauperatum)

I'm always amused by the tiny red cow-udders of cowbag clover (sometimes called poverty clover). I've never observed pollinators visit this diminutive one inch plant, but they delight me nonetheless.

Lithophragma aff. parviflorum

Woodland stars, Lithophragma spp., are small scented flowers in the Saxifragaceae. They are pollinated primarily by invariably dull Greya moths. The moths are usually grey or white, and quite slender.

Olsynium douglasii

Grass widows, Olsynium douglasii, are relatively large beautiful flowers in the Iridaceae (related to Iris, Crocus, Crocosmia, and others). They are often the first blooms found on the top of Upper Table Rock, growing up to twelve inches. Oddly, I've never witnessed bees visiting the flowers despite seeming very attractive. Large bees like bumble bee queens are probably important pollinators for grass widows. An interesting feature of O. douglasii is that nectar is excreted directly from the filaments (stamens). (See Rudall et al. 2003)

Limnanthes aff. floccosa

Meadowfoam, the source of one of my favorite varietal honey, was here and there on the Upper Table Rock summit.

Chickweed monkeyflower (Erythranthe alsinoides)

Monkeyflower likes growing where seeps or springs keep the soil wet. I don't often see bees visit these plants, but they're popular with mason bees (Osmia spp.) where I've surveyed for bees.

Vernal pool

One of the features of both Table Rocks are the many vernal pools, home to tiny freshwater fairy shrimp (Branchinecta lynchi). By summer, the pools are completely dry. The miraculous lifecycles of the minuscule shrimp contend with this by going into a type of dormancy. Mosses and various plants like to grow on the margins of these pools allowing bees and other insects to safely land and drink.

Garter snake (Thamnophis sirtalis)

The vernal pools are hotspots for a variety of wildlife. I was surprised to find a garter snake hidden in the marshy grasses!

Burrow entrance of mountain pocket gopher (Thomomys monticola)

Undoubtedly, garter snakes and other predators hunt the various vermin that call the Table Rocks home. Gophers serve an important role on the mesa, tilling the soil and creating perfect soil beds for seeds to germinate. Many native plant seeds need direct soil contact as well as sunlight to germinate.

Dog tick (Dermacentor variabilis) ♂

With vertebrates come parasites. Ticks are an unfortunate risk when hiking, and it's important to be on the lookout. You can reduce your risk of picking up a tick by staying on the trail and thoroughly checking your clothes and body before you get in your car and when you get home. Ticks are like tanks and can survive for days or even weeks without food or water, even in extreme heat. I once kept a tick in a sealed jar on my dashboard in summer, the thing survived for a week!

Sayfly (Tenthredinidae)

Sawflies are related to bees and wasps, are are probably most similar to the ancestral predecessors of all Hymenoptera. Most sawflies feed on plant material, but adults occasionally feed on carbohydrate-rich nectar.

A view of Mount McLoughlin midway up the Upper Table Rock trail

Upper Table Rock trail is not very long or difficult, but it's worth taking your time to view the surrounding infrastructure surrounding the Table Rocks. Farms and ranches contrast with the nearly pristine landscapes flanking the trail. It's a bit sad.

White leaf manzanita (Arctostaphylos viscida) and honey bee

Manzanita bloom was starting to peak at the time of my hike. The highest diversity of bees could be found on these large shrubs. Honey bees might not be the best equipped to pollinate the flowers or reach the nectar due to the constricted urn-shaped flowers. Poricidal anthers keep pollen restricted only to bees that can sonicate, or buzz pollinate, the flowers. Sonication is when the bee grabs onto the flower and vibrates the flight muscles, a feat honey bees haven't evolved to do.

Habropoda depressa ♂

Mountain digger bees, Habropoda depressa, can and do sonicate flowers. White-leaf manzanita are commonly visited by the bumblebee-sized digger bees.

Nomada aff. erigeronis

Nomad cuckoo bees (Nomada spp.) are beautiful red kleptoparasites of Andrena and other solitary bees. They are in the Apidae which encompasses honey bees (Apis spp.), bumble bees (Bombus spp.), digger bees (Anthophora and Habropoda spp.) and others. Males patrol host nest sites and mimic the odors of the host species, which are deposited on the females during mating. This allows female Nomada to enter host nests without being detected. These fascinating wasp-like bees always make my heart skip a beat. They are so awesome! Not awesome for the host bees, since the cuckoo bee larvae kill the host larvae with large sickle-shaped mandibles, then proceeding to feed on the pollen provisions provided by the host mother.

Andrena

A potential host for the cuckoo bees, mining bees are quite fond of manzanita blooms and are probably the most numerous bee types on the shrubs at any given time. They probably visit the flowers primarily for nectar, and not so much for pollen.

Andrena

Mining bees can't access the tight manzanita flower entrances to reach the nectar, so they are often seen biting holes in the sides of the flowers to rob the nectar. This is very disadvantageous for the plants, but allows the short-tongued bees to feed. Other bees may find the holes in the flowers and exploit them by enlarging them.

Beefly (Bombyliidae)

A variety of other pollinators are attracted to the abundant manzanita blooms. Beeflies are convincing bee mimics which look like flies puffballs with stick legs. They are mostly parasitoids of ground nesting bees (often Andrena and Halictus). Instead of crawling into the nests of their hosts, they hover over the ground and, with a thrust of their butts, jettison their eggs directly into the entrance of their hosts' nests.

Hummingbird

Hummingbirds are also quite fond of the small flowers! Each individual flower probably produces miniscule quantities of nectar for the relatively large floral visitors, but manzanitas make up for it by producing hundreds of thousands of flowers per shrub.

Thrips (Orothrips sp.)

Often viewed as pest insects, thrips are likely to be very important to hummingbirds. I have suspected for many years that hummingbirds are seeking thrips along with the nectar when visiting manzanitas.

Garrya fremontii

Silktassel is one of those shrubs that I've walked past a hundred times before noticing it, but it was quite beautiful in bloom. Garrya is dioecious, having both male and female plants. The one photographed above was a male, offering only pollen. Silktassel is wind-pollinated, which makes sense considering I observed zero pollinators of any kind anywhere near it.

Garrya fremontii flowers

Bees may visit Garrya if nothing else were available, but it is likely the pollen is deficient in nutrients and protein. There are many bees that collect wind-pollinated pollen, even some specialists.

Here is a view of the unnerving dichotomy between Upper Table Rock and the surrounding landscape. There was a time, long before I was born, when everywhere was as serene and beautiful as Upper Table Rock. Humans have a superiority complex.

Children had left touching and important messages for the hikers. Treat wildlife very nice. The message everyone needs to hear. Take heed!

Floral Visitors MMXXI

noreply@blogger.com (Travis Owen) — Wed, 03 Mar 2021 04:23:00 +0000

Halictus exploiting Scilla mischtschenkoana

Today was the first day of this year that I've seen native bees in my garden. Upon reflection, I started gardening for the purpose of feeding bees. Each year I experiment with new plants, different species, and different gardening methods. In all honesty, it's chaos. Variables in nature are infinite, so everything I've learned has mostly been on accident. That's just the way nature works, everything acting on everything else. Most pollinators are pretty adaptive and will exploit any resource that works for them. This is what makes gardening so interesting. It's like rolling the dice and seeing what you get. This is especially true with nonnative garden plants, which is unfortunately most garden plants available.

Scilla mischtschenkoana, with the ambiguous common name "white squill", is the earliest scilla to bloom in my garden. This is the first year I've ever grown it en masse, and I'm incredibly pleased with the results! Being one of the few things in bloom, the warm weather today has enabled bees to gather some nectar and pollen. A small ground nesting bee, about half the size of a honey bee, spent over ten minutes on a small patch. She may have been there longer, but I only watched for ten minutes. While the anthers appear blue, the pollen is actually a yellow buff color.

A honeybee and Scilla

Of course, honey bees were there. Honey bees usually have other bees beat by numbers early in the spring since they go through winter with a large workforce. Bulbs, similarly, also get a head start. The reason many bulbs can bloom so much earlier than their herbaceous cousins is that they've stored energy in the form of a sugary bulb. It's like the little plants had a few Red Bulls early in the morning and are wide awake before the sleepy forbs. By blooming earlier they don't have to compete with as many plants. Come mid spring or early summer and there are many more things in bloom and competition is supercharged.

Crocus vernus

Crocuses along with tulips and daffodils are practically synonymous with spring. Unlike daffodils and tulips, most crocuses are highly attractive to bees and flies. They are also incredibly easy and reliable, growing in both sun and shade. As a rule, most spring bulbs flower earlier in soil warmed by the sun. Soil temperature may be part of what triggers blooming.

Bombus melanopygus

I was super excited to find the years first bumble bee in one of the large Dutch crocuses! As temperatures rise, bumble bee queens begin to emerge from their winter hideouts. Like a bear just coming out of hibernation, they're hungry, so they look for flowers to restore their lipid reserves so they can start a nest and rear the most important brood of the year. These firstborn are tiny versions of workers who help the queens transition from foraging to solely egg-laying machines.

A fly feeds on crocus pollen

Flies are some of the unsung heroes of the pollinator world, often being lumped into the "other pollinators" category. It's not well known that flies as a group are much more diverse than bees, and pollinate plants in environments and climates that aren't favorable for most bees. For instance, flies are probably more important pollinators in alpine and tundra environments where bees are restricted to days with favorable weather. Flies are also active earlier in the year than most bees, thus they are crucial for the pollination of many winter blooming plants.

Oestroid fly feeding on pollen of Merendera sobolifera

A newcomer to my garden, the fun-sized Merendera sobolifera has won the affections of this fly, and my heart. This Merendera has traveled all the way from Eastern Europe to my garden, where its weird finger-like corm was planted in the fall with unknown expectations. I have been delighted to learn it to be one of the earliest bulbs to bloom in my garden, much to the delight of the fly.

The fly spent more than ten minutes (again, I was only there for ten minutes) exploring the multiple blooms for what I can only assume was delectable pollen, to a fly anyway. Many flies relish pollen and nectar alike, though this one seemed particularly keen on the pollen. It even went so far as to dine on pollen that had fallen off of the anthers onto the petals. I can only wonder if flies are this plants primary pollinators in its natural habitat in Eastern Europe.

Many bulbs, however tasty us humans can imagine them to be, are ignored by pollinators in my garden. One such bulb is the spring snowflake, Leucojum spp. They're related to daffodils which similarly don't attract pollinators as well as other spring bulbs. Leucojum flowers aren't particularly showy, but are quite nice in macro.

Another bulb that has been regretfully ignored by bees in my garden is the winter aconite, Eranthis spp. I think as I increase the number of the plants in the garden they will become more enticing to the pollinator force. Their location in the yard may also prove unproductive since it's a shady area, while bees prefer plants in the sun this early in the year.

The witch hazels started blooming sometime mid January, always the very first blooms of the year. Strange cononut-esque shavings pretend to be petals, while the little cups in the middle emit a nice perfume in close proximity. Flies seem to like them, as do I.

More newcomers to the garden are the sweetbox (Sarcococca spp.) shrublets. They'll eventually become respectable in stature, but apparently they'll take their time. Maybe my grandkids will see the full grown shrubs.

Definitely not going to lie, Euphorbia myrsinites is a total nerd plant. Otherworldly scaly serpents with alien flowers. Too cool! This spurge grows in terrible conditions and loves it. Mostly sunny spots with little water. It asks very little but gives much.

Seeing all the newly awakened pollinators and flowers is as encouraging as it is exhilarating. Early spring not only gives me hope, but also anticipation for what is to come! It's only a matter of time until the leafcutter bees come out of their nests.

Grass carrying wasps, Isodontia spp., have also found my bee nest box to meet their needs. Seeing thin waisted wasps flying by carrying pieces of grass several times larger than they are is always somewhat of a surprise. I have yet to pop open one of their nests, but I have read these wasps provision the nests with dozens of orthopterans (usually small katydids and the like).

Speaking of wasps, I saw the years first paper wasp. European paper wasps, Polistes dominula, introduced to the US from the Mediterranean sometime in the 1970s. Unlike other Polistes wasps which prey primarily on caterpillars, these invaders hunt many types of insects. Ants, small flies, and all manner of soft bodied insects are potential prey.

Polistes dominula is troubling for me. While I must accept they are inimical to the native paper wasps, they are beautiful. They are also fascinating, like other paper wasps, and like it or not they are here to stay. These wasps are part of the grand experiment that we have imposed on the earth, and someday we may finally see the results in all their impossible complexity.

Cynipid wasp (possibly) in a snow queen flower

Wasps show up in unexpected places. The tiny-flowered snow queen, Synthyris reniformis, blooms in my garden. Upon close inspection, a tiny insect sheltered inside a flower. Upon closer inspection, this insect appeared to be waspish in nature. What a delight! Lacking better magnification, identification is probably unlikely. Can knowing make it any more charming?

My finger added, for scale

Wonder is a thing I hope I never lose, and something I'd like to encourage in my children. Keep looking down!

I Miss Summer: A Soliloquy

noreply@blogger.com (Travis Owen) — Tue, 09 Feb 2021 19:54:00 +0000

Winter is a time for reflection. Not intentionally, it just happens. In the fall I feel a sense of relief that the weather has begun to cool and the year comes to its inevitable conclusion. There's also a feeling of remorse as nature seems to die and the days become monochrome and bleak. Nature is taking its annual nap, and although there are small signs of life returning to what has seemed a desolate wasteland, I still long for summer. I miss bees, wasps, and flowers. Watching life other than my own unfold in front of my eyes in the garden or in the wild is a welcome break from my excessively important human problems. If you read that in a sarcastic voice, you are correct.

Eriogonum pyrolifolium var. coryphaeum with a mason bee (Osmia sp.)

Color! I miss color. Spring and summer, from a winter perspective anyway, was full of color. Of course in summer I'm complaining about heat. Aren't us humans fickle? Bees and wasps don't worry about such things. I doubt whether insects can worry at all. They're also probably incapable of envy, unlike myself. Insects have significantly fewer neurons than humans, but they're far from unintelligent. How do you even measure intelligence in non humans? Humans are good at doing human things, but we would probably fail at doing everything a bee does with as much efficiency as they have.

Poppies provide no nectar, but produce a plentitude of pollen!

Consider nest building. I can barely manage to keep my apartment clean, but bees make nests without plans while working together, and they keep their nests impeccably tidy. My apartment, on the other hand, is a walk in fire hazard. The word hovel comes to mind. Solitary bees usually build simple nest tunnels, though for them housekeeping is a matter of life and death. Insects have many parasites and their developing young are particularly vulnerable to attacks from an array of diseases and parasites. Fortunately, as a human, we're immune to everything. Except car accidents. And coronavirus. And poor life choices. Oh, never mind.

Small bee, impressive proboscis.

Insect morphology is perfectly adapted to their livelihoods, quite literally. See the tongue (proboscis, actually) on this bee above? Many types of bees are adapted to forage on a specific variety of flower types. Some specialists are adapted to forage on a single species. Humans don't have this kind of specialization. We're more like swiss army knives, able to do a lot of different things but not great (without great effort) at any one thing. The same life-specialization can be observed in all insects. Humans need years of training and practice to become specialized, but it's not built-in.

Wasps are cool. I miss wasps. And, by extension, I miss aphids even though they suck! But they attract some of the most peculiar wasps. There are several aculeate (having stingers) and parasitoid (having ovipositors, egg laying appendages) wasps that prey specifically on aphids, not to mention ladybird beetles, lacewings, and other predatory arthropods which destroy the little suckers in varied fantastical ways. Aculeate aphid predators collect the aphids to provision nests to feed their young. Parasitoids interpolate their eggs directly into the aphids body using an ovipositor, with the larvae feeding on the living and (sometimes) paralyzed host.

Squash bee butts in a squash blossom

Bees, the hippie cousins to wasps, are also pretty darn cool. Growing squash in my garden guarantees squash bees. It was reassuring to look into the male flowers of a squash blossom on a summer morning and seeing cute little squash bee butts. No matter how my week had been, I knew I could count on that. Gardening has been a great outlet for stress and related maladies of my human life. It was and continues to be a great and effective distraction. I wish I had the focus of a bee whose objectives are objectively simple and straightforward. They are literally grown to fulfill the needs of their lives. Humans have too much freedom. Free will sometimes feels like my enemy. In my modern human world I have the option to do nothing and it won't result in my death (not quickly, anyway). If a bee were to do nothing for an afternoon, death would surely result from dehydration or predation.

Bombus bifarius and Orthocarpus cuspidatus on Mt Ashland, fits like a glove!

I've recently been trying to resist the urge to feel resentment towards humanity for the negative effects our species has had towards the environment. There is no denying that destruction of habitat and pollution have been devastating to the natural world and its denizens. There's no question that I contribute to the destruction of nature in various ways, least of which is likely a significant carbon footprint. But we, too, are part of nature. We are apes able to contemplate our own existence and we make things like houses, cell phones, and pencils. Bees make nests out of mud, leaves, and a variety of excretions (like beeswax). These things made by bees are part of nature. By extension, are things make by humans part of nature? A ball point pen may take a lot longer to break down in the soil. We need to start looking for fungi that can eat plastic.

A eumenine wasp (or maybe something else) on Angelica capitellata

I feel like gardening is one way I can give back to nature. Instead of the clean and sterile tea rose garden, or the barren commercial landscapes adorning all the best Walmart Supercenters, I prefer a more feral garden. Fungi freely send mycelium through natural and diverse mulches, plants share the same soil where one cannot discern where one plant ends and the other begins, and insects do as they will with little interference from the gardener. Walk into a forest, and plants know no bounds. And so it is in my garden. In this way, life can do its thing. Life goes on under the mulch throughout the winter, and I find solace in that, despite what appears to be a dead cold landscape.

A weird fly does yoga on Aconitum columbianum

My garden is just a microcosm of the gardens grown in nature. Currently covered in snow, the garden on Mt Ashland is a playground for my mind and soul. I find that this is the only context I ever mention my soul. I don't know what a soul is, but the wilds of the mountains here are good for mine. Words fail to capture what the blooms of summer in the mountains means to me. So I'll stop writing now.

The best thing I could ever do is share nature with my kids. They are my favorite flowers.

Summer Bees, Wasps, and Other Pollinators

noreply@blogger.com (Travis Owen) — Thu, 27 Sep 2018 00:20:00 +0000

Summer in Southern Oregon starts when the rain stops. In the beginning it is mild, warm, and generally pleasant. In recent times the pleasant warm days in the start of summer give way to smoke and excessive heat. Wildfires in the region seem to be the new normal, and the smoke typically lingers for months until the first autumn rains.

A mating pair of European wool carder bees.

Pollinator activity seems to be reduced in the smoky mid and late summer, but does not cease completely. For many bees and wasps summer is not just the only time they are active in the year, but also the time that they seek and find mates. Unlike honey bee queens which do not visit flowers and mate high in the air, most solitary bees and wasps mate among plants and flowers. Males often stick around a particular area that they have made their own territory, and some species fiercely defend this territory from other males by chasing and grappling with other males. This behavior is particularly noticeable among male carpenter bees (Xylocopa spp.) and wool carder bees (Anthidium spp., pictured mating above).

Megachile rotundata visiting a basil flower

Leafcutter bees (Megachile spp.) and related bees (family Megachilidae) are a common sight in summer. They are distinguished from other families by possessing scopae (specialized pollen collecting hairs) on the underside of their abdomens rather on their legs like most bees. Of course, females are the only ones which have scopae since males don't collect pollen or provision nests. In spring this year I released 40 alfalfa leafcutter bees (M. rotundata) in my garden to observe them. Unfortunately, few took a liking to the nesting materials I offered, and their cocoons were under constant attack from ants and earwigs. The ones that survived had a marked liking for the basil I had planted.

Megachile sp.

Many other species of leafcutters live in Southern Oregon, though only one (M. rotundata) is reared intentionally and used for commercial pollination. Due to their characteristic pollen collection method, often passing their abdominal scopae over pollen laden anthers, they usually have a preference for flowers that have exerted anthers, such as many daisy type flowers in the Asteraceae.

Megachile perihirta

Most Megachile nest in tunnels in wood or woody stems, though some nest in the ground. The common name, leafcutter, is a hint at how they create their nests: by lining and partitioning cells with leaf fragments they have chewed from surrounding plants. Leaf matter is adhered together with saliva. Some, I have found, nest in mere crevices since the leaf bits form cell walls that are strong enough to deter some nest parasites.

A leafcutter bee and a paper wasp are seemingly oblivious to each other as they feed on nectar from narrowleaf milkweed

A yellowjacket on elegant tarweed foraging for budworms

Wasps are another familiar sight in the summer. While many wasps are active in spring, colonies of social wasps are more conspicuous since by this time they have grown to large sizes and now forage frequently for food for their developing brood. I was watching a yellowjacket hop from flower to flower one morning, thinking she was foraging for nectar. While many wasps do visit flowers for the sugary food, this one was foraging for caterpillars feeding on the flowers. Like honey bees, yellowjackets remember where to look for favorable foods, and this one had learned to search the tarweed flowers for her bounty.

Aphid wasp (Pemphredoninae) and aphids

This summer I grew a few large evening primrose plants. Aphids seemed to be highly interested in these large plants, and attracted many other interesting insects. Ants fed on the sugary frass, the honeydew, while syrphid flies and ladybugs deposited eggs which hatched out to become aphid eating larvae. My favorite aphid predators were the aphid wasps. These crabronids captured hundreds of aphids per day to provision their nest cells and feed their young.

Sand wasp (Bembix sp.) burrowing into her sandy nest

Another summer favorite, the sand wasps, were a joy to observe. These wasps nest near water on sandy banks. Their nests are in burrows hidden by the loose sand. Each time a female enters or exits her nest, the entrance disappears behind her. They enter their nests by digging furiously with specialized bristles on their front legs. They prey primarily on flies, and provision each individual cell with dozens of paralyzed flies.

Philanthus gibbosus

I am always happy to see the beewolves, genus Philanthus. Sandy riverbanks are a great place to spot them, as they seem to prefer loose sandy soils for their nests. I have encountered two species, P. gibbosus and P. crabroniformis. Markings on the thorax and the color of the eyes can aid in differentiating the two, with the latter having green eyes and the former (pictured) with black eyes.

Philanthus gibbosus on Melilotus alba

Beewolf is a common name referring specifically to the European beewolf (P. triangulum) but freely used to describe all species in the genus. As the name suggests, they prey nearly entirely on their close cousins, bees. American species prey mostly on solitary bees (usually Halictidae) but will occasionally prey on honey bee workers or sometimes even other wasps or (rarely) flies. Unlike many other wasps, their prey is killed, not paralyzed, before being deposited in their nest. Some species feed their young progressively, meaning the mother wasp brings prey to her developing larvae.

Thread waisted wasp nectaring on narrowleaf milkweed

If you love watching pollinators as I do, watching wild milkweeds in bloom never disappoint. Bees, flies, and butterflies are always usual visitors on milkweed flowers, but wasps are the visitors I look for most. Thread waisted wasps (Ammophila spp.) are common flower visitors, but in my experience they are wary of my presence and make observation a challenge. Not this time. This large thread waisted wasp was completely oblivious to me as I took photos. Upon close inspection of the photos, I became aware of the dozen or so pollinia attached to this wasps feet and palps. Milkweeds, like orchids, shed pollen as sticky packets. These sticky pouches containing potentially hundreds of individual pollen grains are then delivered by the wasp to a receptive stamen of another flower.

Ammophila sp.

Thread waisted wasps are kind of a treat for me, and I rarely see more than one at a time. On a hike up Southern Oregon's Mount McLoughlin in April this year (2018) I was fortunate to witness one thread waisted wasp excavating a nest in the sand of the subalpine zone. The wasp first removed the pebble she had placed over the entrance then proceeded to excavate sand particles and deposit them away from the entrance. I didn't witness the completion of the nest, but they normally finish by placing a pebble over the entrance and concealing it with sand (or soil) and nearby debris to disguise it from nest parasites.

Pompilid wasp on wild carrot

Spider wasps are some of my favorite wasps. I've probably said that about every wasp by now, and I've never lied. Spider wasps, family Pompilidae, are special. Have you ever heard the phrase nature is metal? (If you don't know what it means, google it.) Well, pompilid wasps are metal. The commonality among spider wasps is that they all use spiders as hosts for their young. Many, but not all, paralyze the spider before depositing an egg. Some deposit the paralyzed spider in a burrow with an egg adhered on its underside. The larva, once hatched, will feed on the living spider until it is ready to pupate. The largest of these wasps are the tarantula hawks, named for their choice of host. Another commonality worthy of note is that most pompilids have species specific host spiders.

Pompilid wasp on wild carrot

Pompilid wasps also have a habit of flicking their wings rhythmically, possibly seeking attention from possible predators as a warning that they could sting. While the large tarantula hawk wasps are known for their incredibly painful stings, many pompilids are small and readily flee rather than stinging. For most the sting is probably reserved for their spider host. The Pompilidae is a very diverse family when discussing biology and behavior, as they run the gamut from predator and kleptoparasite to parasitoid, depending on the species.

Ichneumon wasp

The true parasitic wasps are differentiated from the bees and stinging wasps by their lack of a stinger. The sting evolved from an egg-laying apparatus called the ovipositor. Parasitic wasps use their ovipositor to lay eggs on or within a host, usually an insect larvae. In the case of ichneumon wasps, the host is usually a caterpillar or less often beetle or sawfly larvae. Ichneumonid larvae develop within the host until pupating and emerging as a fully formed adult. The host is usually dead or dying by the time the wasp is pupating.

Chalcidoid wasp on dodder

Chalcidoid wasps are not closely related to the ichneumonoids, but they exhibit similar parasitism in some cases. They are a diverse and large group, with behavior ranging from gall formers to kleptoparasites. They are also often very small.

A honey bee and bumble bee visiting Phacelia tanacetifolia

And now back to the bees. I wanted to cover wasps first to illustrate the diversity among this group called the Hymenoptera, the order which encompasses bees, wasps, ants, and sawflies. Bees evolved from wasps millions of years ago. Plants coevolved with pollen adapted for insect distribution. To attract bees, many bee-pollinated plants evolved pollen with relatively high levels of proteins compared to pollen from wind-pollinated plants. There are a few species of bees which don't collect or eat pollen as a protein source (Trigona spp., not found in the US) and a few stinging wasps which do collect pollen as a protein source (Masarinae, vouchered in Oregon though I've not yet encountered them).

Long-horned bee on Madia elegans

Most bees native to the US live solitary lives and don't make honey. Long-horned bees are related to honey bees, but rather than living in a colony nest in congregations of underground tunnels. Each female creates her own tunnel and provisions her own brood chambers. Most solitary ground nesting bees, like most solitary bees in general, create a ball of pollen with a small amount of nectar to hold it together, lay an egg on it, and seal the chamber. Most never interact with their offspring (like many solitary wasps) and only live for a matter of weeks.

Cuckoo bee, Sphecodes

Some of the most peculiar bees in Oregon are the cuckoo bees. They don't collect pollen or make nests. Instead, females seek out nests of other types of solitary bees and lay their eggs in the host brood chambers. The cuckoo larvae kills or starves the host larvae, then develops normally to adulthood. I view cuckoo bees as a sign of a healthy host population of bees.

Ceratina on a wild Campanula

Small carpenter bees are very numerous in my area. They collect very little pollen on their bodies, collecting it instead in their crop and regurgitating it once back to the nest. Nests are built in dried pithy stems of woody plants, linear, with partitions built from the chewed pith.

Ceratina on knapweed

Ceratina visit a wide range of plants in a wide range of families. Their small size allows them to climb deep inside flowers with nectar that is inaccessible to larger bees.

Halictus on chamomile

The Halictidae, sometimes called sweat bees, are some of the most common bees in summer in some areas. They have bushy scopae, specialized pollen collecting hairs, and a notch at the tip of their abdomen (a good diagnostic feature).

Augochlora on cutleaf blackberry

Metallic green bees of the Halictidae are a special treat for the photographer.

Masked bee, Hylaeus, on wild carrot

Like Ceratina, masked bees collect pollen inside a crop and regurgitate it in the nest. These are very small bees and are often found visiting flowers in abundance in the summer, but their small size means they are probably overlooked most of the time.

Large carpenter bees, Xylocopa, on evening primrose

Some of the largest bees in the US, and my garden, are the large carpenter bees. They are in the minority of bees which excavate tunnels in wood to create their nests. They seem to prefer dead wood, such as the planks of my fence, and unpainted decks and siding of homes around the country. These large, docile bees visit a wide variety of plants over a long season. They rear two to three generation a year in my yard. Occasionally females overwinter in their nests and interact with their offspring, but I suspect this is the exception rather than the rule. Due to their ability to generate their own warmth by vibrating their flight muscles isometrically they can fly later in the day. In the middle of summer they were flying long after the sun fell to forage on the evening primrose (primarily night blooming).

A white-lined sphinx moth, highly attracted to the highly perfumed evening primrose

Pollinators and Wildflowers of Mount McLoughlin

noreply@blogger.com (Travis Owen) — Thu, 19 Jul 2018 06:41:00 +0000

Mount McLoughlin is one of the most stunning and recognizable natural landmarks in the Rogue Valley. With an elevation of 9,493ft it is the sixth highest mountain in Oregon. It is a volcano, and the entire mountain is littered with volcanic rocks. The mountain is located between Medford and Klamath Falls in the Sky Lakes Wilderness. But all that aside, I have looked on at Mount McLoughlin with wonder. I wondered, what is up there? What is it like to be on that mountain? I'm not an experienced hiker, but I had to do this.

Dicentra formosa (Fumariaceae) in the subalpine zone in June

I made four separate treks on the mountain, but only made it to the summit on the most recent hike. My first climb was August 2017. My decision to experience the mountain was a spur of the moment choice. I started on the trail later than was needed to make the summit and descend before darkness fell, so I descended before summiting. My second attempt was in November 2017, one day after another hike which I had achieved a minor injury. The cold and my injury were too much, so I gave up early on the trail.

A view from the ridge, high winds and clouds enveloped the top of the mountain.

June this year I was back on the mountain. Little was in bloom, and it was fairly cold. I had arduously reached the ridge, but dark clouds were moving in around the mountain from the East. Thunder was rumbling from the distance. Before I knew what was happening, the storm was right overhead. Snow and lightening prompted me to make a swift descent! In my slightly panicked haste, I lost the trail several times on the way down through the subalpine zone (dominated by many large basalt fragments, dwarf trees, and a poorly discernable trail). It was not ideal.

Wasp-mimic syrphid fly (Chrysotoxum) eating pollen on Lilium pardalinum

I encountered much more pleasant weather on the mountain this July. Driving up to the trailhead on a pitted rocky dirt road, I passed by so many wildflowers in bloom! It was exactly what I needed to boost my spirits and give me the optimism I needed to reach the summit, finally.

Pacific coralroot orchid (Corallorhiza mertensiana; Orchidaceae)

Gleefully on my way up the trail, eyes mostly to the ground, I spotted a delightfully intriguing plant! Of course I encountered many interesting plants, but one that stood out among the rest was the parasitic coralroot. Instead of photosynthesizing, these orchids get all they need from relationships with ectomycorrhizal fungi. The fungi gets what it needs to live from relationships with the roots of surrounding conifers.

Pompilid wasps

As I progressed up the trail, I noticed the characteristic wing flicking of spider wasps (Pompilidae). Four or five small pompilids were very interested in a small area on the side of the trail, maybe a few inches in diameter. Most pompilids hunt spiders to paralyze and feed to their young, but some are kleptoparasites of other pompilids. I found one such kleptoparasitic pompilid last year. I can't help but wonder if these too were kleptoparasites attracted to the hidden burrow of another pompilid.

Scree on the southern face of Mount McLoughlin

Finally, and arduously, I reached the subalpine zone. The trees are sparsely distributed, dwarfed, and contorted. Soil, if you could call it that, is highly mineral, mostly sand and fine gravel. Wildflowers were blooming everywhere in pockets between large rocks. Deep roots keep the plants anchored in the loose dry substrate. Hiking at this stage becomes challenging, and the trail is frought with many obstacles. This is when the fun begins.

Eriogonum pyrolifolium (Polygonaceae)

Life high on the mountain is hard for plants. Prolonged snow cover, harsh weather, strong wind, loose soils, a short growing season, and animal predation creates a very trying environment for high elevation plants. Most of these plants cope by having deep extensive root systems and small above ground manifestations. Many bloom very quickly as soon as weather permits to cope with the short growing season. Summer for us in the lowlands is spring on the mountain.

Pinus contorta in the subalpine zone

The trail becomes slightly harder to follow as one gets higher on the mountain. It's not that there isn't a discernable trail, but that there are many discernable trails, likely from hikers that came before you that went the wrong way. My view is that it is hard to get lost going up. All you have to do is just go up. Going off the beaten path might make the ascent more difficult, as was my case. This was partly due to my desire to A.) see more wildflowers, B.) explore my own limitations, and C.) avoid other hikers.

Bombus mixtus on Eriogonum marifolium ♀

During my hike, the most numerous wildflowers were a few species of Eriogonum, or wild buckwheats, namely E. marifolium and E. pyrolifolium. The yellow flowered E. marifolium is interesting in that it is dioecious, that is, plants are either male or female. Female phenotypes have flowers which turn red as they age, while male phenotypes have all yellow flowers with a shorter corolla. In my observations, eriogonums are visited by a wide range of pollinators.

Large pompilid wasp on Eriogonum marifolium ♂

While there were bees present on many of the mountain wildflowers, the most conspicuous floral visitors were spider wasps. Large and small pompilids were on nearly every eriogonum I encountered. In other words, sit and watch an eriogonum and within a minute there would be at least one, sometimes two, pompilids on the flowers.

Pompilid wasp visiting Eriogonum pyrolifolium

Spider wasps are some of my favorite wasps. Not only do I think their way of life is brutal and cold (AKA very cool!) but they have a beautiful form. While they are rarely brightly colored, they do attract attention by their rhythmic wing flicks and jolty movements. Some species are known to have painful stings, something I would like to explore in greater detail.

Eriogonum pyrolifolium

Eriogonum pyrolifolium has an odd common name: dirty socks. The internet claims it is a tip of the hat to their delightful smell. I haven't thought to give them a whiff. Yet. Maybe the next time I see these flowers I will remember to smell them.

Bombus vandykei queen and worker on Penstemon davidsonii

Davidson's penstemon is found in varying elevations on the mountain, normally in rock crevices. It is probably exclusively visited by bees. I have not seen any pollinator but various bees visit it on my various hikes up the mountain.

Bombus on Castilleja arachnoidea (Orobanchaceae)

Paintbrush are strange plants. They are thought to be parasitic on the roots of other plants growing nearby. Flowers are hidden within bracts that must be folded back by bees to access the nectar. Osmiine bees and bumble bees were foraging on them during my hike, and interestingly enough, it was the first time I've seen any pollinator visiting Castilleja.

Ammophila nest entrance, covered by a single pebble.

Thread waisted wasps (Ammophila; Sphecidae) are pretty awesome, and plentiful on Mount McLoughlin. They are fairly nervous creatures. Their nests, which are provisioned with paralyzed caterpillars, are constructed in sandy soils. They have the peculiar behavior of using a pebble to cover the entrance while they are looking for prey. When the nest is provisioned and eggs are laid, the entrance is covered completely by sand and debris to hide any sign of a nest to discourage parasites.

Ammophila female above her nest entrance

By a stroke of luck, I happened to be sitting and drinking some water on the scree when a thread waisted wasp alighted right next to me. She removed the pebble covering the entrance to her nest (which I hadn't noticed previously) and proceeded to continue excavation.

When thread waisted wasps are excavating their nest tunnels, the excavated sand is deposited away from the entrance of the nest so as not to draw attention to the location of the nest entrance.

Luetkea pectinata (Rosaceae)

Partridgefoot (Luetkea pectinata) is a plant that seems to grow exclusively on the south facing scree. I haven't ventured onto the northern or western slopes to determine if it grows there, too. Flies sem to have been the primary pollinators, though bumble bees were also seen on the flowers.

A view down the northeastern flank

Did I mention it was hot? It was hot. I was staying hydrated, but a pounding headache was making the already difficult trek even more difficult. I started to feel like Frodo on Mount Doom trying to deliver the One Ring to the firey pit from whence it came. At this point my entire body was jelly and I was questioning my life choices.

Hulsea nana (Asteraceae)

While eriogonums still managed to hold some ground in the alpine zone (there are no trees here), Hulsea nana was the champion of the harsh conditions of the top. The exquisite foliage alone make this plant a treat, and the bright yellow daisies were encouraging. Small flies were visiting the flowers, but I had lost the will to wait for a good photo opportunity. Onwards and upwards, keep going up! (I kept joking to myself that there were just a few vertical miles to go.)

Eriogonum pyrolifolium near the summit with... a honey bee?!

Yeah. I get within ten minutes of the summit and I see a honey bee. Why did I not see a single honey bee anywhere else on the mountain? Where did she come from?! I wonder if she was from a managed hive somewhere at the base of the mountain, or a feral colony hidden within a rock crevice or tree hollow. The mystery will haunt me until I die.

A tiny wasp lands on me once I reached the summit, Platygastridae?

I have long since unlearned the habit of immediately brushing off an insect that finds its way onto my skin. First, I look to see what it is. In this case, it was a tiny wasp! The quality of the photo is probably too poor to ever get a positive ID, but knowing that wasps are equally attracted to me as I am to them makes my heart happy.

Horntail (Urocerus; Siricidae) on the Mount McLoughlin summit

So, I made it to the summit. It was epic! There were exactly zero plants growing on the tip top, but there must have been thousands upon thousands of insects; mostly flies and horntails. Woodwasps, sawflies, and horntails are ancient phytophagous (plant feeding) progenitors of wasps. Siricid woodwasp larvae feed on decaying wood of conifers.

A view of fellow hikers on the summit, looking southeast

More Bees from Spring '18

noreply@blogger.com (Travis Owen) — Sat, 23 Jun 2018 05:02:00 +0000

Here we are halfway through June. How is it already summer? It seems like just a few weeks ago the pears were blooming and I was working in rain and cold trying desperately to get the spring work done with the honey bee hives. Now it feels like summer is in full effect, rain is less and less in the forecast, and multitudes of bees are busily seeking mates and provisioning nests!

Cuckoo bee (Nomada, Apidae) on arrowleaf buckwheat (Eriogonum compositum, Polygonaceae) in the wilds

The most familiar and recognized bees are honey bees, poster child of pollinator conservation alongside monarch butterflies. But the overwhelming majority of native bees in the US and the world are solitary and nearly half of those collect pollen from only one family of flowering plants. Of course, the plant family varies between different bees. Many of the most common bees are generalists who visit a wide range of flowers from many families. In either case, a diverse range of flower types from a range of families is the best for facilitating a diversity of bees.

Male carpenter bee (Xylocopa tabaniformis, Apidae) on comfrey (Symphytum officinale, Boraginaceae) in my garden

Large carpenter bees, Xylocopa spp., are very conspicuous in my garden. My yard is fenced with ratty looking cedar boards serving as habitat for countless Xylocopa nests. The boards are only about 3/4" thick, but the large bees somehow create vertical nests within the boards without breaching the surface. I think this is incredible, and I have no idea how they can detect the outside edge of the wood.

Carpenter bee on a foxglove (Digitalis purpurea, Plantaginaceae) in my garden

Foxgloves, native to this part of Oregon, grow in my garden. Carpenter bees have no idea how to get in, so instead they just bite holes at the top. Even then, they are not very attractive to most of the bees in my garden.

Male carpenter bee on wood spurge (Euphorbia amygdaloides, Euphorbiaceae) in my garden

Wood spurge, Euphorbia amygdaloides, was surprisingly attractive to some pollinators in my garden. Carpenter bees, beetles, flies, and even paper wasps were attractive to the odd musky smelling flowers. Unlike most flowers visited by bees, euphorbia flowers don't produce nectar. For the species that do produce nectar, it is secreted from bracts below the actual flowers, sometimes appearing like petals or sepals which are absent from all euphorbias.

Xylocopa, a mosaic gynandromorph, perched on a Salvia (Lamiaceae) in my garden

Speaking of bizarre, did you know that some hymenopterans have both male and female characteristics? This is called gynandromorphism. Sexual dimorphism, present in the Hymenoptera, is where sexes of the same species exhibit differing characteristics beyond their genitalia (i.e. scopae, not found on male bees). I stumbled upon one or possibly two Xylocopa gynandromorphs in my garden.

Xylocopa, a mosaic gynandromorph, perched on a Salvia (Lamiaceae) in my garden

I took photos of the carpenter bee, found in the morning before it had warmed up enough to fly, and hadn't realized it might be a gynandromorph until looking at the photos later. It was a mosaic gynandromorph, only having patches of male characteristics rather than being a bilateral gynandromorph (literally half male and half female). Web searches revealed that gynandromorphism isn't unheard of in Xylocopa, but maybe not all that common either.

Xylocopa gynandromorph under magnification (microscope courtesy Old Sol Bees). Male (L) and female (R)

After sharing the images of my Xylocopa gynandromorph on social media, I felt regretful for not having captured the insect. Returning home, I immediately went to my garden, net in hand, and watched as the carpenter bees foraged for resources and males chasing each other off. I captured and released a few males and one large female. I was about to give up when I saw a peculiar individual. I netted it exiting a nest tunnel, and sure enough it was a mosaic gynandromorph. It was an intermediate size between a typical male and female, and had scopae on both legs. It appeared to be provisioning a nest, so perhaps it was also fertile. Gynandromorphs are generally considered to be infertile. The specimen has been vouchered, and I am interested in donating it to a museum.

Halictus (Halictidae) visiting a foxglove, correctly, in my garden

As mentioned previously, few bees visited the foxgloves in my garden. The only bees I observed visiting the flowers properly (sans nectar thievery) were small carpenter bees (Ceratina) and sweat bees (Halictus). The most numerous visitors to the foxgloves were ants. They were probably attracted to the nectar, but ant pollination in this case seems unlikely because of how the reproductive structures are positioned within the flowers.

Halictus on a buttercup (Ranunculus sp., Ranunculaceae) in the wilds

Halictus is one of the most common genera that I observe. They visit a wide range of flower types, but mostly open flowers with either easily accessible pollen or nectar. Many can be easily identified by the small notch seen at the tip of the abdomen. Buttercup pollen is toxic to some bees, or at least to their offspring, though it seems to have no effect on some bees.

Small bees foraging on deerbrush (Ceanothus integerrimus, Rhamnaceae) in the wilds

Deerbrush (Ceanothus integerrimus) and the earlier blooming buckbrush (C. cuneatus) are both very attractive plants to many pollinators as well as this human. They are shrubs, and when in bloom are covered in panicles of tiny simple flowers. Most of the pollinators are also very small, typically bees and beetles. They are graceful and rewarding plants which also fix nitrogen into the soil with specialized root nodes formed by a bacterial partner.

Halictus ligatus on a yellow yarrow (Achillea sp., Asteraceae) at the Jackson County OSU Extension in Central Point, OR

When I first started researching what to plant for bees years ago, yarrow was one of the first plants I tried to grow. It was fairly easy from seed, and took practically no water once established. Yet, over the years I observed few pollinators visiting the flowers. Ladybugs are probably the most frequent visitors that I can recall. This yellow variety seemed to garner the attention of a variety of small sweat bees (Halictidae) and small carpenter bees (Ceratina) at the OSU Extension in Jackson County.

Halictus ligatus foraging on narrow leaf mule ears (Wyethia angustifolia, Asteraceae) in the wilds

A larger and more popular relative of yarrow, narrow leaf mule ears are very popular with bees where they bloom in forest clearings around the Rogue River area. Many types of bees visit Wyethia, including bees from the families Apidae (including honey bees), Halictidae, and Megachilidae.

Osmiine bee (Megachilidae) visiting narrow leaf mule ears

I attempted W. angustifolia from wild-collected seed. I managed to get the seeds to germinate with cold stratification (a fancy way to say I left the pot outside through winter), but killed most off after that. There were a few small individuals which I successfully managed to not kill into their third year, but they died promptly after transplanting into my garden. Perhaps they are not currently in cultivation due to tragically fragile root systems, or I just need to do a better job next time.

A honey bee foraging on carrotleaf horkelia (Horkelia daucifolia, Rosaceae) in the wilds

One of my favorite plants, perhaps only due to obscurity, is an uncommon rose relative native only to Southern Oregon and Northern California. The plant is somewhat uncommon, so when I happen upon it I feel fortunate and grateful. Pollinators of all types visit it for nectar and pollen, the open flowers don't care who is pollinating it. Many types of bees, flies, beetles, and butterflies are always attracted to it in my observations, weather permitting. Where situated near an apiary, it attracts honey bees.

Honey bees love trailing snowberry (Symphoricarpos hesperius, Caprifoliaceae) in the wilds

Snowberry, like carrotleaf horkelia, is very unassuming. Creeping or trailing snowberry likes growing in somewhat shady locations under a canopy of conifers or in clearings. The flowers, small and easy to miss, are typically hidden under the foliage. Honey bees don't have any trouble finding the flowers. Snowberry honey is highly sought after, apparently, though the author has never tried it. I have only ever seen honey bees visit these flowers, though I would bet they are also popular among bumble bees since not many bees forage in shady locations besides bumble bees.

European woolcarder bee (Anthidium manicatum, Megachilidae) forages on lamb's ear (Stachys byzantina, Lamiaceae) in my garden

Back in the garden, lambs ear is in bloom with fuzzy flower spikes interspersed with small purple flowers. European woolcarder bees, an introduced species, have a special relationship with this plant and seem to be attracted to it wherever it is grown. Females forage almost exclusively on the lambs ear, while males fiercely defend the area from other males and most other bees by chasing or grappling.

European woolcarder bee (Anthidium manicatum, Megachilidae) forages on lamb's ear (Stachys byzantina, Lamiaceae) in my garden

Woolcarder bees, introduced or native, are in the Megachilidae. All megachilid females collect pollen on scopae under their abdomens rather than their legs like most other bees. Woolcarder bee females also collect hairs from plants such as lambs ear to line and partition cells in small tunnels and other small cavities.

Small carpenter bee (Ceratina, Apidae) on a Penstemon in the Jackson County OSU Extension in Central Point, OR

Small carpenter bees, very unlike their larger relatives, nest in dried pithy stems of plants like roses or many woody perennials. These bees are very small, easily mistaken for small flies, but are also very common. Here I witnessed one perched on a penstemon flower to groom itself. The penstemon flowers are deep and narrow inside, perfect for small bees to climb right in to reach the nectar and in doing so brushing against the reproductive organs.

Cuckoo bee (Nomada, Apidae) on arrowleaf buckwheat (Eriogonum compositum, Polygonaceae) in the wilds

Growing in hot dry rocky locations by the Rogue River, arrowleaf buckwheat attracts a range of tiny flies, beetles, butterflies, wasps, and bees. One peculiar bee, one of my favorites, was visiting the eriogonum during one of my photo sessions. This is a nomad cuckoo bee related not too closely to honey bees and carpenter bees. The cuckoos are kleptoparasites, nest thieves, of other ground nesting bees.

Cuckoo bee (Nomada, Apidae) on arrowleaf buckwheat (Eriogonum compositum, Polygonaceae) in the wilds

As nest thieves, cuckoos require neither to collect pollen nor to build a nest. Needless to say, they don't possess specialized pollen collecting hairs, so they often appear very wasplike. Females fly low to the ground in a zigzag pattern trying to detect the scent of a host nest. Once a nest is located, she will enter and deposit one of her eggs in a finished or nearly finished cell, sealing it behind her. The cuckoo larvae either kills or starves the host larvae, and develops on the host provisions.

Masked bee (Hylaeus, Colletidae) on arrowleaf buckwheat (Eriogonum compositum, Polygonaceae) in the wilds

Yellow faced bees, or masked bees, are also wasplike due to lacking pollen collecting hairs. Instead of collecting pollen on scopae or corbiculae like other pollen collecting bees, they eat it. The pollen in regurgitated from their crop when they return to the nest, and formed into a ball within each cell.

Masked bee (Hylaeus, Colletidae) on arrowleaf buckwheat (Eriogonum compositum, Polygonaceae) in the wilds

Because cuckoo bees and masked bees do not have much hair on their bodies, they are not thought to be very effective pollinators. However, some pollen adheres via a static charge or with microscopic velcro-like projections. Flying insects, especially bees, develop a positive charge from flight while grounded objects like plants carry a negative charge. This helps the pollen become unstuck from the anther and attracted to the insect. The impact of the charge on the pollen is minimal, but every bit counts!

See the "mask" of a male masked bee

Bees, among other fascinating insects (like wasps), are fun to learn about! Observing which flowers they visit and what type of bee is visiting is an ongoing experiment that everyone can take part in. Gardens present an additional dimension of interest. Watching native pollinators visit nonnative garden plants or weeds, or watching nonnative bees visit native plants, is an interesting way to study the theory of pollination syndromes. I imagine someday someone could develop a pollination syndrome chart for specific types of bees based on flower shape, color, characteristics of pollen or nectar, et cetera. Maybe such a chart could aid in conservation efforts geared towards at-risk species of bees or other pollinators. Until then, just go watch some flowers and see what comes of it.

Bees in Spring 2018

noreply@blogger.com (Travis Owen) — Mon, 14 May 2018 07:15:00 +0000

Spring is the best time in Southern Oregon to watch bees! The combination of highly diverse wildflower communities, excellent climatic conditions, and a relative abundance of water culminates to produce a great situation for many species of bees. Without looking up the data, I would bet that the highest diversity of bees can be found in the spring months in Southern Oregon. Not only have I seen more different kinds of bees in the spring than in summer or fall, but there are more native plants in bloom in Southern Oregon than later in the year which offer a diverse nutritional palate.

A honey bee clings to a willow catkin with one leg while grooming the pollen into her pollen baskets. February 22nd.

The earliest blooms in Southern Oregon are found on the trees. Alder, hazel, and willow are some of the first trees to bloom and produce an abundance of pollen for the few bees active at this time as well as a plethora of flies and beetles that feed on pollen. When apiaries or the odd feral honey bee colony are located near any of these trees, often near bodies of water, they delight in the abundant pollen on warm sunny days. Few native bees are active in January or February when the earliest blossoms appear, but I saw Andrena and a few bumble bee queens foraging on willow catkins in February this year.

A mating pair of Ceratina on pear blossoms. April.

Southern Oregon in April is host to billions of pear blossoms. Pear orchards are fairly commonplace around Medford, though many have been sold and turned into housing developments or hemp farms, much to the dismay of bees and other pollinators. Pear growers use honey bees to pollinate their trees, but the beekeepers make a fraction of what they are paid per hive in the almond orchards in California. Where almond growers may pay up to $200 per honey bee hive in the almond orchards of the Central Valley in California, pear growers rent less hives and might pay up to $40 per hive. This is probably because almonds bloom much earlier than pears, a time when few native pollinators are active, and almonds are currently much more valuable than pears. Consider the cost of pears compared to the cost of almonds next time you go grocery shopping.

A mating pair of Ceratina on pear blossoms

The pear orchards of Southern Oregon, at least the ones I have been in, are surrounded by a lot of natural landscapes and filled with flowering weeds. Whereas almonds are harvested by shaking the trees and then raked from the ground, the floor of the almond orchards needs to be bare or mowed. Pears are not collected this way, so the ground beneath the trees is sometimes overgrown with weeds. This offers a lot of resources for native and managed bees alike.

Osmia on a crab apple blossom. May.

Apple blossoms follow the pears. I am not sure there are many apple orchards in Southern Oregon, but they are grown as ornamental trees and personal orchards. Mason bees, Osmia spp., are sometimes managed for pollination of spring tree crops including almonds. Unlike honey bees, each female creates her own nest and does not live socially. In fact, most female native bees fulfill the role of both queen and worker. Osmia are called mason bees because they use mud to build the cell walls and partition cells in their nests (preexisting linear tunnels, usually in wood).

Nomada on a crab apple blossom

Peculiar, wasp-like, and fascinating. That is how I like to introduce the cuckoo bees. As the common name implies, they steal the nests of other bees. Unlike the birds of the similar name, cuckoo bees lay an egg in the cell of a host bee, such as Andrena or Halictus, and the cuckoo bee larvae usually kills the host larvae and develops on the food (a ball of pollen) collected by the host bee. Since the cuckoo bees steal nests from other bees, they don't create nests themselves. Since the nest they steal is provisioned by the host, they have no need to collect pollen, and thus they don't have pollen collecting hairs on their bodies.

Nomada female searching for host bee burrows

Cuckoo bees in the genus Nomada are usually black, yellow, red, or a combination of those colors. I was lucky to see a few red and black females hovering just inches from the ground recently. They were seeking host nests, probably Andrena or Halictus which nest in soil. Nomada females find Andrena nests by olfactory cues (Cane 1983).

Anthophora female on Phacelia tanacetifolia. May.

Phacelia tanacetifolia, or lacy phacelia, is a member of the Boraginaceae, native to California and possibly Oregon. It is more commonly available as seed and usually grown as a cover crop, ornamental, or sometimes specifically for bee forage. It doesn't let down, it is a great bee plant. Bees of all shapes and sizes, as well as a variety of other types of pollinators, are attracted to the blue scorpioid inflorescences.

Anthophora female on Phacelia tanacetifolia

I recently had the opportunity to observe bees visiting a large planting of P. tanacetifolia at Easy Valley Farm in Rogue River, OR. One of the more conspicuous types of bees were the large and fast digger bees. Anthophora is a common and widespread genus with some species remaining active from spring to fall. While some solitary bees only have a single generation each year, some species have two or multiple generations. At least some if not most species of Anthophora produce offspring all year, with the last generation of the year overwintering as prepupae within underground nests.

Halictus on Phacelia tanacetifolia

Sweat bees, more accurately known as halictid bees since not all species are attracted to sweat, are very active in spring and summer. They vary widely in size and nesting habits, and are very widespread and speciose.

Bombus vosnesenskii clinging to an Anchusa azurea blossom

An Old World member of the Boraginaceae, Anchusa azurea is found in my garden and in those fortunate enough to have acquired it from seed. It is like a large forget-me-not, and is very attractive to bumble bees and honey bees. I photographed an odd flower with only four lobes, most have five, with a bumble bee clinging to it early one morning. This bumble either was caught in the cold and unable to return home, or wanted an early start.

Iris chrysophylla with a bumble bee

Iris chrysophylla, yellowleaf iris, is a dryland iris found among grasses in mixed coniferous forest clearings and edges. It blooms profusely in mid spring over a long period in Southern Oregon. Various bees visit the flowers, including mason bees, halictid bees, and bumble bees. Though it is visited by various bees, I do not typically see a lot of bees visiting the flowers even where it is blooming in large masses.

Bombus diving for nectar on Iris chrysophylla

Iris flowers, as well as a few other Iridaceae relatives found around the world (i.e. Hermodactylus, Moraea), are complex and peculiar. To oversimplify, each single flower is divided into three ports of access for bees to enter seeking nectar. Pollen is typically deposited onto the back of the bee, but only if it enters the correct way. Bumble bees have very long tongues and are able to circumvent the reproductive structures and access the nectar from the side.

A honey bee forages for nectar on Calochortus tolmiei

While Iris chrysophylla grows from a short rhizome, Calochortus tolmiei grows from a bulb. Calochortus is only found in the New World and is very unlike other New World bulbs in flower structure. C. tolmiei is one of a handful of species with peculiar hairs on the petals, likely an adaptation to prevent inefficient pollinators from stealing the nectar. Nectar is excreted from colored patches at the base of the petals, and the hairs force the bee to rub against the anthers to access it.

Calochortus tolmiei and a honey bee fleeing from a camera that came too close for comfort.

Arctostaphylos viscida

noreply@blogger.com (Travis Owen) — Sun, 25 Mar 2018 01:35:00 +0000

From left: Callophyrs augustinus and an empidid fly nectaring on Arctostaphylos viscida.

Arctostaphylos viscida Parry (Sticky whiteleaf manzanita) is in full bloom here in Southerstern Oregon. Manzanitas are in the family Ericaceae, which includes related genera Arbutus (madrone, strawberry tree), Calluna (heather), Erica (heath), Kalmiopsis, Pieris, Rhododendron (includes azaleas), Vaccinium (blueberries, cranberries, etc.), and over a hundred more genera, few of which known in cultivation. A. viscida is native from Southwestern Oregon south through most of California (to Kern County, possibly further south). It is a shrub or small tree, sometimes reaching a height of fifteen feet in optimal conditions, shorter further south. Small panicle-like clusters of urn-shaped flowers appear in Spring. The entire inflorescence is sticky, covered in glandular hairs, with glaucus leaves, thus the common name.

Arctostaphylos viscida, a variant with white florets

Sticky whiteleaf manzanita grows in association with oak (Quercus), Pacific madrone (Arbutus menziesii), various conifers (Juniperus, Pinus, Pseudotsuga), and Ceanothus among others. With many of these, it shares a variety of endo- and ectomycorrhizae, which are types of fungi which survive on or within the roots of a host and trade valuable nutrients (as well as increasing the surface area of the roots) for carbon and other exudates (root secretions). Dozens of species of mycorrhiza have been discovered in association with Arctostaphylos and associated plant species. Essentially, mycorrhizal fungi are symbiants which infect their hosts, but they benefit the plants in multiple ways and are necessary for the establishment and survival of countless taxa in the plant kingdom. Like all ericaceous plants, Arctostaphylos requires symbiosis with specialized ericoid mycorrhizal fungi to survive in the acidic and nutrient poor soils they are typically found in.

Arctostaphylos viscida flanks the northeastern rim of Upper Table Rock in Central Point, OR

Arctostaphylos viscida is a pioneer shrub, entering the landscape in succession following grasses and other herbaceous plants. A. viscida paves the way for trees such as Douglas-fir (Pseudotsuga menziesii), and contributes to its establishment by hosting the necessary mycorrhizal fungi that Douglas-fir requires to survive. The same group of mycorrhiza also aids in the establishment of Arbutus menziesii, a relationship which may explain why madrone is so difficult to keep alive in cultivation (lacking the correct mycorrhizal symbiosis outside its native range).

Flowers of Arctostaphylos viscida with banded thrips, Aeolothrips sp.

The flowers of Arctostaphylos viscida are similar to many (but not all) ericaceous plants such as Arbutus, Pieris, and Vaccinium by having fused petals forming an urn or cup, a stigma nearly twice the length of the stamen, and a ring of stamen surrounding the ovary and nectaries. Manzanita pollen is dehisced through small pores and often requires sonication, or buzz pollination, to release the pollen. Small insects such as thrips, which are small enough to crawl into the flowers, can gather pollen without the use of sonication. A. viscida may occasionally be self-compatible but progeny probably benefits most from cross pollination between different plants. Bloom time varies between individual specimens, as does the relative attractiveness to pollinators due to unknown factors that may include the health of the plant, quality of the nectar, or something else entirely.

Arctostaphylos viscida bark

The bark is similar to that of madrone (Arbutus menziesii): cinnamony, thin, and smooth once the outer layer of bark peels off. It has the unique characteristic of repelling water. This feature may contribute to the plants ability to survive in severe drought, though if there is inadequate rain or sun they do not flower well. Arctostaphylos viscida shrubs that grow on the north side of structures or under dense canopies of trees do not seem to flower reliably. The excessive shade from a northern exposure or tall trees promotes fungal infections that result in aborted buds and black spots on the leaves. A. viscida prefers open sunny habitats, and is more fit in chaparral than in dense woodlands, though it does perfectly well at forest edges, clearings, and slopes which in some aspects have similar characteristics to true chaparral.

From left: A. viscida berries, crushed dried berries, and seeds.

Seeds of Arctostaphylos are produced in the form of berries. In A. viscida the berries are sometimes sticky, but this seems to go away when the fruit is mature. There is speculation that the berries (and seeds) are distributed by animals, probably birds. Laboratory tests of seed procured from coyote feces (what a fun job that must have been) found low germ rates. Birds are more likely, as the berries would likely pass through quickly (acid treatment in a lab helped break seed dormancy). Fire is another treatment that helps scarify the seed and break dormancy, and has the added benefit of reducing competition (especially tree saplings) which would otherwise shade out the slow growing manzanita. Suppressing fire is probably hindering the development of new colonies of manzanita. By briefly examining many of the stands in my area, it is difficult to find small young plants, but mature plants 6' or taller are common.

A honey bee, Apis mellifera, seeking nectar

Many insects, including the introduced European honey bee (Apis mellifera), visit manzanita flowers where they are abundant. An individual plant can produce hundreds of thousands of individual florets that open over the span of a few weeks. Bees and flies are the most common visitors to the flowers, though the abundance of either depends largely on the surrounding habitat. For example, plants growing among conifer forests may have more flies than bees, since conifer forests tend to support less bees than do chaparral or oak woodlands. Honey bees will work the flowers when both are abundant, though in dry years they may not be able to reach the nectar since most plants produce less nectar in drought. Other insects occasionally bite holes in the sides of the flowers to reach the nectar, and when that happens honey bees will gain access to the nectar through the side as well though it is unlikely they will create the holes themselves.

A bumblebee queen, Bombus vosnesenskii, collecting nectar.

The primary pollinators of manzanitas are typically medium and large bees which are capable of sonication, or buzz pollination. The bee lands on the flower and vibrates the flight muscles which releases a burst of pollen from the pores in the anthers. A positive electric charge (which all flighted insects generate) helps the pollen cling to the bee. Bumble bees (Bombus), mason bees (Osmia), digger bees (Anthophorini), and long-horned bees (Eucerini) are capable of sonication and I have observed all visiting A. viscida in my observations.

Male Andrena sp., robbing nectar through a hole in the side of the flower.

Many smaller bees visit the flowers, and in my observations many of them are nectar thieves. It's unclear who is chewing the holes in the flowers, but once they are there the flower has a low chance of getting pollinated since most of the visitors to those flowers bypass the reproductive structures and feed on nectar through the hole.

A cuckoo bee, Nomada sp., robbing nectar from a hole in the floret.

I have personally observed many types of small bees visiting A. viscida including mining bees (Andrena), small carpenter bees (Ceratina), and cuckoo bees (Nomada). Cuckoo bees are kleptoparasitoids, like cuckoo birds, who lay their eggs in the nests of other bees. Cuckoo bees forgo the creation of their own nests, and do not collect pollen. Instead, cuckoos rely on the provisions supplied by host species, usually other closely related bees. The invader sneaks into the nest while the host is out and lays an egg in a cell chamber. When the cuckoo bee larvae hatches, it kills the host larvae then feeds on the host provisions and develops in the host nest until adulthood. The genus Nomada, sometimes called nomad bees, are in the Apidae, the family that includes honey bees, bumble bees, digger bees, carpenter bees, and a few more. Mining bees, Andrena, are typical hosts for nomads, but others include Agepostemon (green metallic sweat bees) and a few others.

From left: a mason bee, Osmia sp., perched on a leaf, and a paper wasp.

Many types of bees mate on and around wildflower patches, and this includes manzanitas in full bloom. I have captured and observed male mason bees (Osmia), male long-horned bees (Eucerini), male mining bees (Andrena), and male carpenter bees (Ceratina) in close proximity to A. viscida. Many mate while foraging, unlike honey bees. While honey bee queens mate with several males and don't forage, the majority of native bees mate with a single male while foraging. Male honey bees, or drones, mate with a single queen and then die, while the majority of native bee males mate as much as possible with as many females as they can mount.

Mischocyttarus flavitarsis

Vespid wasps are typically just getting started while A. viscida is in bloom in Southern Oregon. Females which hibernated through winter are just beginning to emerge pending good weather, and they are keen on acquiring the much needed carbohydrates supplied by flowers in the form of sugary nectar. The beginning of the year for yellowjackets and paper wasps is not dissimilar, as mated females must create their nests and feed the first of their offspring before matured workers are able to take over the foraging responsibilities so the queen can stay home and devote the remainder of her life to laying eggs. Yellowjackets, genus Vespula, are similar to honey bees in that there are distinct castes (reproductive queens, males, and workers). If yellowjacket workers do lay eggs, they will only rear male offspring since the workers are not equipped to mate. Paper wasps (Polistes and Mischocyttarus) have females that are all capable of mating and thus laying both male and female offspring. While there are no true paper wasp queens, a dominant laying female asserts her sovereignty by fighting with competing females (size is often a factor of success).

A parasitoid wasp, Ichneumonini

Many parasitoid wasps congregate on and around A. viscida for nectar and honeydew, as well as to seek out hosts. I've observed ichneumonoids and chalcidoids either seeking hosts or seeking sustenance. Many suitable hosts are found on A. viscida, including many lepidopteran caterpillars.

Male ant, Lasius sp., stuck to the sticky inflorescence.

The sticky inflorescences of A. viscida trap many small insects, notably small flies and winged ants. Winged ants were one of the unfortunate insects to get stuck to the inflorescence. Ants in the genus Lasius are often attracted to honeydew, the frass of sap sucking insects like scale or aphids. Subterranean species of Lasius feed on the honeydew of root feeding aphids, forming mutualistic relationships.

A bee fly, Bombylius major, feeding on nectar

Flies visit A. viscida flowers frequently, and may also be gathering around the plants to mate or seek out prey (predatory species). More diverse and speciose than bees, flies are reliable pollinators in climatic conditions not accommodating to most bees. Many flies mimic bees and wasps, although flies are incapable of stinging. Bee flies, Bombyliidae, not only do a good job at mimicking bees, some are even parasitoids of ground nesting bees.

Empididae

Dagger flies, or dance flies, are named for the long rigid proboscises they possess and the distinctive courting rituals performed by the males. Not all species have a long proboscis, but those that do use them to probe flowers for nectar. I have seen them visiting many early blooming trees such as Prunus and Salix, and early blooming ephemerals like Crocus and Muscari. They are also predatory, feeding on smaller flies, while larvae feed on decaying matter in the soil.

Empididae infected with Entomophthora fungi

I've seen several dead flies clinging to leaves and flowers, having been infected with the fungi Entomophthora. Spores are forcibly ejected from specialized structured grown by the fungus from membranous sections of the abdomen. Flies under or downwind may inadvertently catch a spore which proceeds to grow into the unsuspecting fly. The fungal hyphae gradually grow throughout the entire body, digesting the flies organs, and when it reaches the brain causes the fly to land and climb upwards. When the fly dies, after about a week from becoming infected, spores are produced and the cycle is renewed. Entomophthora isn't restricted to flies, some species infect aphids, mites, grasshoppers, and others.

Noctuidae caterpillar

A handful of moths in the Noctuidae and Geometridae use A. viscida as a host plant. Caterpillars are often difficult to see unless one is seeking them out, due to their drab colors and patterns. Adult moths of various types visit the flowers for nectar.

Nymphalis californica

During warm weather, butterflies visit the flowers. I have found this to be true only in clearings with a lot of direct sunlight. Manzanitas that receive only filtered light appear to be less attractive to butterflies. Adult butterflies were seen sunbathing as well as nectaring, while on a few occasions males attempted to defend their territory from other species. I suspect courtship may also occur in the vicinity as well, though I didn't observe it directly. California tortoiseshells, Nymphalis californica, overwinter as adults and benefit from access to early nectar sources such as A. viscida.

A mayfly, Ephemeroptera

Mayflies live for about a day as adults, and are incapable of eating. Thus, a mayflies presence on A. viscida flowers was purely coincidental. Mayflies spend one to several years as aquatic nymphs before reaching adulthood, feeding on algae and detritus from under rocks at the bottom of streams, rivers, and occasionally lakes.

Banded thrips, Aeolothrips sp.

Thrips, at least in some areas, are very numerous on and within the flowers of A. viscida. Thrips feed primarily on plant tissue, including flowers, buds, and pollen. But despite their reputation, they may also be pollinators. Thrips have been demonstrated to be pollinators of many plants in many regions of the world, from tropical environments to high altitude and high latitude alpine and tundra landscapes, respectively. They may also be some of the most ancient pollinators. Whether or not they benefit A. viscida through direct pollination or not, they may serve the plant by attracting hummingbirds which feed on them as well as feeding on nectar, facilitating pollination. Rufous and Anna's hummingbirds, Selasphorus rufus and Calypte anna, respectively, are suspected to feed on both thrips and nectar during their visits. Conversely, high numbers of thrips on a particular inflorescence appear to keep other pollinators (i.e. bees) from visiting that inflorescence in my observations, although the thrips occupation varies from inflorescence to inflorescence on a single plant.

References:

Acsai, Jan, and David L. Largent. "Fungi Associated with Arbutus Menziesii, Arctostaphylos Manzanita, and Arctostaphylos Uva-ursi in Central and Northern California." Mycologia 75.3 (1983): 544.http://goo.gl/x2fXFD

Armstrong, W. P. "Termite, Winged Ant, Small Wasp or Gnat?" Ant Genera Index. Wayne's Word, 2013. Web. http://waynesword.palomar.edu/AntGenera.htm.http://goo.gl/rlkVld

"Calflora: Arctostaphylos Viscida." Calflora: Information on California Plants for Education, Research and Conservation, with Data Contributed by Public and Private Institutions and Individuals, including the Consortium of California Herbaria. 2016.http://goo.gl/vbJaA

Clarke, D., H. Whitney, G. Sutton, and D. Robert. "Detection and Learning of Floral Electric Fields by Bumblebees." Science 340.6128 (2013): 66-69. Web.https://goo.gl/T6P02x

Eliyahu, Dorit, Andrew C. McCall, Marina Lauck, and Ana Trakhtenbrot. "Florivory and Nectar-robbing Perforations in Flowers of Pointleaf Manzanita Arctostaphylos Pungens (Ericaceae) and Their Effects on Plant Reproductive Success." Arthropod-Plant Interactions 9.6 (2015): 613-22. Web.

Eliyahu, Dorit, McCall C. Andrew, Lauck Marina, Ana Trakhenbrot, and Judith L. Bronstein. "Minute Pollinators: The Role of Thrips (Thysanoptera) as Pollinators of Pointleaf Manzanita, Arctostaphylos Pungens (Ericaceae)." J Pollinat Ecol. 16 (2015): 64-71. Web.http://goo.gl/aghCgK

Fryer, Janet L. 2015. Arctostaphylos viscida, sticky whiteleaf manzanita. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Missoula Fire Sciences Laboratory (Producer).http://goo.gl/IDAtp4

Horton, Thomas R., Thomas D. Bruns, and V. Thomas Parker. "Ectomycorrhizal Fungi Associated with Arctostaphylos Contribute to Pseudotsuga Menziesii Establishment." Can. J. Bot. Canadian Journal of Botany 77.1 (1999): 93-102.http://goo.gl/Wzdt0G

Lewis, Donald. "Why Do Some Ants Have Wings?" ISU Extension News Release. ISU, 22 July 2002. Web.http://goo.gl/7AUR7V

Moncada, Roberto. "Pollinators and Nectar Robbers on Manzanita (Arctostaphylos Sp.)." Amateur Biology Blog. 07 Feb. 2013. Web.http://goo.gl/ZtRAvw

Rostás, Michael, and Jürgen Tautz. "Ants as Pollinators of Plants and the Role of Floral Scents." All Flesh Is Grass Cellular Origin, Life in Extreme Habitats and Astrobiology (2010): 149-61. Webhttp://goo.gl/KXDd5t

Shirley, C. 2011. "Plant Guide for sticky whiteleaf manzanita (Arctostaphylos viscida)". USDA-Natural Resources Conservation Service, California Plant Materials Center, Lockeford, CA 95237http://goo.gl/dckZS5

Turner, Mark. "Arctostaphylos Viscida | Sticky Whiteleaf Manzanita." Wildflowers of the Pacific Northwest. Web.http://goo.gl/ekPEz2

Parasitoid Wasps of Southern Oregon 2017

noreply@blogger.com (Travis Owen) — Tue, 13 Feb 2018 02:30:00 +0000

Anyone that knows me knows that I love wasps. I think you should love them, too. Here I will attempt to familiarize you with the world of the non-stinging wasps known as the parasitoids. Parasitic wasps do not have true stings, as the aculeate wasps [and bees] do. These parasitoids have ovipositors, which are used to lay, or sometimes inject, eggs. While there are aculeate parasitoids, the aculeates do not have ovipositors. (The exception is the Chrysididae, the cuckoo wasps, which are aculeates which evolved their own unique ovipositors independently from the parasitoids featured in this piece.) The aculeate sting evolved from an ovipositor many millions of years ago.

Female snakefly with ovipositor, Raphidioptera

Wasps are in the super diverse order Hymenoptera, which also includes bees, ants, and sawflies (including horntails). This huge, and old, lineage holds more species than there are known vertebrates in the world. The orders Hymenoptera, Coleoptera (beetles), and Raphidioptera (snakeflies), among a few other orders, are thought to have evolved from a common ancestor in the Permian period. The earliest hymenopterans, the sawflies, suborder Symphyta, first appear in the Triassic period. Most sawflies (and horntails) are phytophagous, meaning the larvae feed on plant material. The first true wasps appear in the mid-Jurassic period, and the earliest known bees in the Cretaceous along with flowering plants.

A colorful male Ichneumonidae metasoma

There are over 130,000 species of wasps and bees worldwide with new species being described each year. Parasitic wasps, more ancient and possibly more speciose than the aculeate wasps and bees, have very diverse features and characteristics not found among aculeates. These features represent the diverse behaviors and life styles of the thousands of parasitic wasps found throughout the world.

Ichneumonidae antennae with many flagellomeres

Sexual dimorphism is common among the Hymenoptera and there are many features that can be used to tell the sex of individuals. The antennae of aculeates have similar numbers of flagellomeres (antennal segments) between families and can be sexed based on the number (females have ten flagellomeres while males have eleven). Parasitoids occasionally, however, have many flagellomeres and the number differs significantly between species. Males don’t lay eggs, so individuals of species which have long ovipositors can be sexed based on the presence or absence of the ovipositor. Males, in general, are difficult to identify to species since many species resemble each other, while the females bear traits that offer the best means to identify a species.

Torymus female, Torymidae, reared from a mossy rose gall

The ovipositor of parasitoids can be long or short, and in many cases is at least partially retractable. In the case of many the ovipositor is protected within a sheath which is usually visible at all times. Lay observers of the unenlightened kind often mistake the ovipositor sheath for a sting and demonstrate unnecessary uneasiness towards the poor creature. Long ovipositors are not adapted to penetrating human skin, nor do the wasps possess a venom that would inflict any pain worth calling mother about. Some species do have short, rigid ovipositors which could potentially poke through the skin if the wasp is handled. The best advice is to leave the wasp alone if at all possible. Even if it is not capable of stinging, your attempt to coerce it onto your face would probably stress it out, and nobody needs that.

Cryptini female

Parasitoid wasps possess venom, though unlike their eusocial aculeate cousins the venom is adapted specifically for host species. Similarly to many aculeate wasps, parasitoid venom is composed of a cocktail of proteins, peptides (including kinins, found in many wasp venoms), and many toxins (the chemistry, admittedly, is beyond my education level). Some of the wasps in the families Braconidae, Ichneumonidae, and Pteromalidae possess polydnaviruses (which evolved independently in all three cases) which serve to interfere with the hosts’ immune system. Venoms injected alongside the polydnaviruses, at least in the case of the bracoviruses (the polydnaviruses possessed by braconids), further aid the pathogen to do its job. In the case of eggs injected into the host, the venoms and polydnaviruses help ensure the survival of the wasp egg(s) and larvae.

Toryminae female; Torymidae, reared from a mossy rose gall

Parasitoid venoms exhibit other characteristics including antimicrobial and biopesticidal properties. The lethality or paralyzing effects are sometimes highly dependent on the age of the host, and in some cases highly species specific. Some venoms send the host into premature pupation which the wasp larvae may need to develop properly. Other venoms simply block neurotransmitters or neuromuscular transmission which may protect an egg being deposited to the exterior of a host. Another effect of the venom of at least some braconids is the alteration of the host hemolymph for the nutritional benefit of the wasp larvae, or in other cases causing the host to become lethargic resulting in easier oviposition or protection for the immature wasps.

Gasteruption and a small solitary bee

Many adult parasitoids visit flowers for nectar or occasionally for sugary sap or honeydew, and the presence of flowers may even extend the lives of adult wasps. Many parasitoid wasps have short tongues and so only benefit from small flowers with very exposed nectar. Flowers in the Apiaceae such as herbs including parsley or coriander as well as vegetables like celery or carrots are all great plants for parasitoid wasps and will also benefit small bees and a myriad of other small beneficial insects. The composite family, Asteraceae, also has many good species for wasps. When choosing species for parasitoid wasps, select species with many small flowers instead of few large flowers. Deep tubular flowers will not benefit short-tongued wasps, but will instead be more attractive to birds, butterflies, or large bees. Planting anything is better than nothing, and planting native plants is best. Parasitoid wasps may indirectly benefit from planting host plants for insect hosts (i.e. plants which attract aphids for aphidiine wasps).

Ichneumonidae

I've been observing wasps in close detail for the last several years. They just fascinate me. When I was a kid I loved watching the silicon-based extraterrestrial parasitoids in the Alien movies designed by the late H.R. Giger, creatures which I still find both terrifying and beautiful. We are fortunate that parasitoid wasps, which aren't too different from the parasitoid aliens in some of their lifestyles, are comparatively small and do not use our bodies as hosts. Below are many of the parasitoids I had observed in 2017 through both direct organic observation and capture. Most of the encounters were completely by chance, but by realizing that these wasps are practically everywhere and knowing what to look for, you too will begin to discover tiny aliens in your backyard, workplace, and probably everywhere else you go.

Cryptinae female with a US cent coin

Ichneumonoidea

Some of the most recognized and speciose of the parasitoids are the wasps in the Ichneumonoidea, the superfamily which includes the ichneumons and braconids. There are over 6,000 species described north of Mexico, and at over 100,000 worldwide. Both ichneumons and braconids utilize a diversity of hosts across many insect orders and even some arachnids in various life stages from egg to adult.

The most peculiar feature among some of the Ichneumonoidea (notably the subfamilies Campopleginae and Banchinae in the Ichneumonidae, and subfamilies Microgastrinae, Miracinae, Cheloninae, Adeliinae, Cardiochilinae, Khoikhoiinae, and Mendesellinae in the Braconidae) is the presence of polydnaviruses in their genomes. Polydnaviruses are released into the host when an ichneumonoid is ovipositing, serving to suppress the immune system of the host and thus protecting the egg. The polydnavirus then infects the developing wasp larvae. Specific polydnaviruses have coevolved with the wasps they infect, and the wasps have evolved to be dependant on the virus to protect the developing wasp larvae.

Ichneumonidae

The Ichneumonidae, or ichneumon wasps, are ecto- or endoparasitoids (external and internal parasitoids, respectively) of a wide variety of insects and arachnids (including immature or adult spiders, and egg sacs of spiders and pseudoscorpions). Their hosts are always insects which exhibit complete metamorphosis (where an insect goes through four distinct stages: egg, larvae, pupae, and adult) rather than incomplete or gradual metamorphosis.

Fused cells in ichneumon forewings create the "horsehead" often used in determining Ichneumonidae from Braconidae. The small hairs, or setae, protect the membranes, help create lift, and demonstrate hydrophobic effects.

Ichneumoninae

The subfamily Ichneumoninae is made up of both ecto- and endoparasitic wasps which parasitize caterpillars of both lepidopterans (mostly moths) and symphytans (sawflies). They have short ovipositors, so must find hosts that are in exposed locations such as external plant feeding caterpillars. Ichneumonine wasps are idiobionts, meaning their hosts cease or slow development after oviposition. Hosts are parasitized typically in the larval or pupal stage, but wasps always emerge from pupae. Adults occasionally eat large portions of the host bodies, sometimes killing them in the process. Adults also feed on honeydew from aphids or other sap sucking insects, and are said to eat the foliage of certain plants themselves.

A male ichneumon (Ichneumonini tribe) which I captured in a field of drying grasses. I captured (and released) many of these individuals in the same field. It is likely they were seeking females to mate with. Male ichneumons are often colorful while females tend to be drab.

Many of the ichneumons are cryptic and only known from host records, meaning a researcher collected infected insect larvae or pupae and watched what emerged from its dead or dying body. Courtship and mating among ichneumons is studied to some extent but few broad scale studies have been done, with most research focusing instead on one to few closely related species.

Captured male ichneumon (Ichneumonini tribe)

Solitary females of most hymenopterans, with the exception of honey bees, usually mate with a single male while males may mate with several females. This is partially true with ichneumons. Solitary ichneumons, that is species which lay a single egg in each host, typically mate with one or less often two males. Gregarious species, females which lay more than one egg within a host, mate with multiple males.

Ichneumonini tribe

Males of most species emerge prior to the emergence of the females, occasionally congregating near the pre-emergent females. Males of some species, such as giant ichneumons, Megarhyssa spp. (Rhyssinae), inseminate pre-emergent females by inserting their elongated abdomens into the exit burrows. Insemination of this type is usually successful, though post-emergent fertilization on the tree bark with freshly emerged females has also been observed although it is less successful.

Early spring in Corvalis, Oregon, I captured and released this ichneumon (Ichneumonini tribe) which was near the entrance of a honey bee hive. The weather was overcast, windy, and wet, and it is possible this wasp was attracted to the warmth of the colony or an associated lepidopteran host.

Male ichneumons seek females with the aid of pheromones produces by the female, or by honing into the scent of a particular host plant (such as a plant in/on which the wasps’ host insect is found). In some cases both virgin and mated females exude pheromones. When male ichneumons discover a mate, they probe the female with their antennae to orient themselves for fertilization. In the case of Pimpla turionellae (a European species), and perhaps many others, male pheromones excreted from specialized glands on the antennae aid in female receptiveness when the males probe the female antennae. This antennal probing is sometimes done throughout the duration of intercourse which can take from between 1 to 20 minutes depending on the species, though mating acts have rarely been directly observed.

Some ichneumon females are ready to oviposit almost immediately after mating. Adult longevity and host availability are inherently connected as statistically the longer an adult female lives the more likely she will find hosts to oviposit in. Floral hosts (i.e. dandelions) and sources of honeydew (sugary frass from sap-sucking insects such as aphids) have been demonstrated in a few economically important ichneumons to increase adult longevity thus increasing their effectiveness as biocontrol agents.

Banchinae

The Banchinae is a fairly common group of parasitoids with 1,500 species worldwide, and 600 north of Mexico. They are diurnal endoparasitoids of various larval Lepidoptera, Coleoptera, and Hymenoptera. Along with the Campopleginae, each species harbors its own genetically unique polydnavirus.

Exetastes (Banchinae)

Exetastes is a fairly common genus found worldwide with the exception of Antarctica and Australia. They inhabit mostly unforested landscapes like chaparral and oak savannas, or sometimes in large forest clearings. Adults visit flowers frequently. I have observed Exetastes visiting Daucus (carrot) for nectar in early autumn. The individual photographed here was captured in low grasses midday on a sunny southern facing hillside.

Exetastes (Banchinae)

Hosts for Exetastes are usually cutworms (Noctuidae). Host larvae live to initiate pupation (whether it is initiated prematurely or not is unknown) in the soil by forming a pupal case. The wasp larvae kills the host after the pupal case is constructed and pupates within it.

Cryptinae

The Cryptinae is the largest family of the Ichneumonidae with over 400 genera worldwide. They are mostly ectoparasites of a range of hosts including lepidopterans, coleopterans, dipterans, and even other hymenopterans. Some are hyperparasitoids of other parasitoids in the Braconidae and other ichneumonids. Females of some species are wingless (i.e. some Gelis).

Cryptinae

Some cryptines are at least partially nocturnal, and drawn to lights on warm evenings. Most are comparatively small in the tribes Claseini (the smallest tribe with only two genera in South America), Hemigastrini, and Phygadeuontini while the largest tribe, Criptini, includes wasps of larger average size for the subfamily.

Cryptinae

Cryptine wasps were pretty common sights for me in the summer of 2017. I had observed many small species, perhaps the same species, in and around a large apiary. The specimen photographed above was found within empty bee equipment in storage inhabited by a few paper wasps Polistes aurifer and the nonnative P. dominula. Whether there was a true association or their proximity was merely a coincidence is left for future discovery. Most cryptines seek out hosts found in soil or in leaf litter, with the exception of the Cryptini which typically seek hosts in foliage.

Acroricnus stylator (tribe Cryptini; Cryptinae)

As previously mentioned, some cryptines use other hymenopterans as hosts. Acroricnus stylator is one such species that seeks out wasp hosts which build mud nests. This includes eumenine wasps such as Ancistrocerus and Eumenes, mud daubers like Sceliphron , and mason bees (Osmia)

Acroricnus stylator (Cryptinae), an individual I caught and later released.

Acroricnus stylator is found in both the New and Old World with various regionally specific subspecies. Adults are known to feed on flowers of the carrot and daisy families. Though they parasitize other beneficial hymenopterans, their presence would suggest healthy host populations.

Xoridinae

Xoridine wasps, such as one I discovered on a door under a porch light on a cool late spring morning, are usually hosts of wood boring larvae of beetles (Cerambycidae) and horntails (Siricidae). Some may use other hosts in the Lepidoptera or Hymenoptera, or may be facultatively hyperparasitic.

Braconidae

Braconids and Ichneumonids are usually differentiated by their wing venation, but some other factors come into consideration. Braconids utilize a wide variety of hosts, including aphids, bark beetles, and caterpillars. Many species parasitize eggs of their hosts, but don't fully develop until the host reaches the larval stage. Unlike the Ichneumonidae, most braconids pupate in silk-like cocoons outside the bodies of their hosts either directly attached to the host or removed from it completely. Also unlike the ichneumonids, few braconids use hosts in the pupal stage to reach maturity, with the exception of the subfamilies Alysiinae and Opiinae.

Meteorus (Braconidae)

Many braconids, such as Meteorus, are economically important to both farmers and gardeners since hosts of these wasps include significant crop pests such as cutworms and tomato hornworms. A few studies have demonstrated that flowering plants increased the lifespans of females, possibly translating into more caterpillars taken as hosts.

Meteorus (Braconidae)

Braconid venom, at least for some species, causes apoptosis (the natural death of cells) around the oviposition site. This could serve to protect the egg, and perhaps protect the young larvae as it begins to feed.

Microgastrinae male captured around stored honey bee boxes infested with greater wax moths

In the summer of 2017, I discovered empty honey bee boxes infested with wax moths (Galleria mellonella) and numerous small cocoons. There were also pteromalid wasps, which I now suspect to be hyperparasitoids of the braconids. The cocoons were likely from microgastrine braconids (possibly Cotesia spp., a common genus). Microgastrines are koinobiont endoparasitoids, mostly on moth larvae. Some species are gregarious, meaning females lay many eggs in a host, while others are solitary.

Silk cocoons of Microgastrinae

Evanioidea

The Evanioidea contains three families: Aulacidae, Evaniidae, and Gasteruptiidae. The Aulacidae are parasitoids of wood boring beetles, mostly Cerambycidae, and some Xiphydriidae (Symphyta). The Evaniidae, or ensign wasps, are parasitoids of cockroach egg cases.

Gasteruptiidae

The family Gasteruptiidae have unique and easily identifiable abdomens. They are sometimes known as carrot wasps, probably a hint of the forage preferences of adult wasps seeking nectar, though they will visit other plants with small flowers that have easily accessible floral resources. For instance, I have observed them visiting flowers of onions, Allium cepa.

Gasteruption on parsley flowers in late spring

Larvae of gasteruptiid wasps are predators or predator-inquilines of other hymenopterans that nest in pithy stems or twigs of woody plants. Predator-inquilines eat the larval food in the nest rather than the larvae itself, while predatory gasteruptiids feed on the larvae itself, such is the case for Gasteruption. Either way, the host larvae dies. Gasteruptiids go through winter as prepupae, then pupate in the spring.

Cynipoidea

Gall wasps, Cynipoidea, are a unique group of parasitoids because most feed on plant tissue. Some are gall inquilines, those that live within the galls of other Cynipoidea though don't create galls themselves. Families Figitidae and Ibaliidae are parasitoids or hyperparasitoids of other insects. Figitidae wasps are parasitoids of various Diptera, Neuroptera, or are parasitoids of various gall inducing wasps in the Cynipoidea and Chalcidoidea (i.e. Cynipidae, Eurytomidae). Some Figitidae are hyperparasitoids of Braconidae or Chalcidoidea parasitoids of Hemiptera hosts. The Ibaliidae are parasitoids of sawflies in the family Siricidae. Early Ibaliidae instar larvae are endoparasitic, but later instars emerge and eat the rest of the host from the outside. The Cynipidae is made up entirely of gall inducing wasps and gall inquilines, and nearly entirely phytophagous (feed on plant tissue).

Trichoteras tubifaciens (Cynipidae), crystalline tube gall. This is my best guess as to the identity of this group of small galls. There are very few resources citing this species, which appears to be restricted to a small distribution from Northern California to Southern Oregon.

Cynipidae

The family Cynipidae is an oddity in the world of wasps (though if you have read this far, it has probably come to light that wasps, particularly the parasitoids, are an oddity as a whole). Cynipid wasps are mostly gall forming, phytophagous (herbivorous) wasps which induce galls in which to feed on and live within during their development. Inquilines don't make galls themselves, but feed on the gall of another wasp, occasionally killing the host larvae.

Galls on the underside of Quercus garryana leaves, likely Cynips mirabilis (the larger speckled gall) and possibly Neuroterus (the small disk-shaped galls).

Gall forming wasps initiate the formation of a gall when certain chemicals excreted by the wasps (or on the egg) make contact with the plant tissue, though the exact mechanism is unclear in most cases. The type of plant or tree as well as the location of the gall is very specific to the species of wasp. Most gall wasps only form galls on a select few or single species on plant or tree. Usually oaks (Quercus spp.) and other trees of the family Fagaceae, or roses (Rosa spp.) are most often used as host plants for gall formation. There are some herbaceous gall formers who favor plants in the families Asteraceae, Lamiaceae, and Papaveraceae, though their galls are less conspicuous.

Large conspicuous galls, like those of Andricus quercuscalifornicus, are subject to predation from birds. Many galls are at least partially hidden from plain view, or may otherwise be camouflaged in some way.

The gall serves two primary purposes for the wasps: as a source of food for developing larvae and to protect the immature wasp from its natural enemies. The inside of the gall is full of nutritious plant tissue, and a source of protein for the wasp larvae. The outside of the gall is often hardened to form a shell-like exterior. Sometimes the outer layers are toxic, a protective measure that keeps predators from eating the gall.

Cynipid galls with emerged Torymus tubicola (Torymidae). Galls are subject not only to predation, but to parasitism by other insects including other hymenopterans.

There are at least fourteen hundred described species of cynipid wasps in the world, 750 in around fifty genera in North America. Many have interesting sex lives. Some species are believed to be completely asexual with completely female populations. Others have strange reproductive cycles. Some bivoltine species (two generations per year) have generations which alternate between sexual and asexual (parthenogenetic), such that the sexual generation which has both male and female members will yield only females. The asexual (entirely female) generation will not mate, yet will yield both male and female offspring. Quite alien when compared to our human lives!

Mossy rose gall, Diplolepis rosae

As the specific epithet implies, Diplolepis rosae forms galls on a select few species of roses. D. rosae is a European native, accidentally introduced into the United States. Wasps lay eggs on the young leaf buds in spring as new leaves emerge. They are gregarious, and multiple larvae feed on a single gall, and they remain inside the gall throughout the winter before pupating and emerging as adults in spring. They are possibly parthenogenetic in some populations, as very few males have ever been documented. Ichneumonid and chalcidoid wasps have been known to parasitize the larvae of D. rosae.

Chalcidoidea

Torymus tubicola, a parasitoid of cynipid wasp galls.

The smallest wasps, and smallest insects, are in the families Mymaridae and Trichogrammatidae, both contain egg parasitoids of very small host insects: Thysanoptera (thrips) and Psocoptera (barklice). The most peculiar species, however, are those which are hyperparasitic and use other parasitoids as hosts. This can be in the form of secondary or tertiary parasitoids, or to the extreme as quaternary parasitoids. This equates to a parasitoid parasitizing a parasitoid parasitizing a parasitoid of some sort of unfortunate host. It is fathomable that such an interaction may even save the original host, but only if the secondary parasitoid parasitizes the primary parasitoid before pupation, the point at which the original host would be severely weakened by the primary parasitoid. Crystal clear, right?

A very small pteromalid on the tip of a decking screw under high magnification

Some chalcidoids are polyembrionic. Polyembriony in wasps is where a single fertilized egg produces more than one larvae, and some will even produce thousands of offspring within a single host. Copidosoma floridanum (Encyrtidae) typically produces broods exceeding 2000 larvae in a single noctunid host. Some of the siblings will be nonreproductive "soldiers" who patrol the hosts body seeking out competing parasitoid larvae and killing them. Polyembriony is not restricted to the Chalcidoidea, but is also found in a few species in the Braconidae, Platygasteridae, and Dryinidae.

Minimal venation is typical of chalcidoid wasps. This is the wing of a pteromalid under magnificatipon.

In all, chalcidoid wasps attack insects in around 340 families representing fifteen orders, as well as some arachnids (i.e. pseudoscorpions, ticks, and mites) including other hymenopterans. Mostly immature stages of hosts are parasitized, but adult hosts are sometimes utilized as well as eggs and egg sacs (including spider egg sacs, in some cases).

Torymidae

Torymids are small, often metallic green parasitoids (occasionally phytophagous) wasps sometimes confused with chrysidids. Most are ectoparasitoids or occasionally hyperparasitoids of gall forming hymenopterans (usually Cynipidae), dipterans (Cecidomyiidae), and scale insects (Coccoidea). Species that live in the galls of others may be inquilines, parasitoids, or both entomophagous and phytophagous by feeding on both the gall and the larvae within (similar to some Eurytomidae). Some gall inquilines don't feed on the host larvae, but kill it anyway then proceed to feed on the gall tissue. A few are seed feeders, a unique group which oviposits within developing seeds where the larvae will develop on the tissue within.

Torymus male which emerged from a mossy rose gall, Diplolepis rosae

I have reared, albeit unintentionally, a handful of torymids from cynipid galls of various types. Galls of Cynips mirabilis, a solitary species, have yielded a single torymid per gall. Mossy rose galls, formed by the gregarious species Diplolepis rosae, yielded several torymids of varying sizes and sexes.

Torymus male which emerged from a mossy rose gall, Diplolepis rosae

Pteromalidae

The pteromalid wasps are mostly ectoparasitoids of Lepidoptera, Coleoptera, and to a lesser extent Diptera and other Hymenoptera (Braconidae). Some are also endoparasitoids or hyperparasitoids, though few if any are phytophagous. Adults only live for a short time, but will feed on nectar, honeydew, or sugary plant secretions. Females require protein for egg production, and host feeding is common. Females typically feed at secretions from the oviposition site, and may even consume some of the host tissue itself. Some species have been described feeding on the eggs of scale insects, as well as serving as biocontrol agents.

Pteromalidae reared from a greater wax moth, Galleria mellonella, under magnification

Pteromalid eggs on a severely weakened greater wax moth larvae

In summer of 2017 I observed and captured pteromalids roaming among greater wax moth (Galleria mellonella) larvae in stored bee boxes. The wax moths had destroyed a great amount of the stored honey bee comb, but there were also many dead moth larvae and Microgastrinae (Braconidae) cocoons. To explore further, I collected several wax moth cocoons to dissect. One contained a severely weakened moth larvae with a dozen small eggs adhered to its body. I placed the larvae in a lidded jar in a temperature controlled room to observe. In two weeks small pupae were developing on the exterior of the larvae with no cocoons of their own, suggesting they rely on the moths own pupal case to protect them. A week later there were adult pteromalids walking around within the jar.

Diaprioidea

The Diaprioidea is composed of the endoparasitoids of various Diptera, and occasionally Coleoptera or other Hymenoptera. They are small, obscure wasps, and only the most basic information is known about their biology.

Diapriidae

At least three hundred species are found north of Mexico, with four thousand species estimated worldwide. They are parasitoids of fungus gnats and other dipterans, and often found in wooded areas with decaying vegetation (where their hosts are common).

Diapriidae with a US quarter dollar

I've observed diapriid wasps on two occasions, both at porch lights in the summer. It may only be a coincidence that their host insects are dipterans, the most numerous insects to frequent my porch lights.

Conclusion

This concludes my observations of parasitoid wasps from 2017. If there is anything I would like you to take from this it is that nature is amazing. And what can you do to protect these obscure yet fascinating creatures? Plant flowers, and try to reduce your use of pesticides. It's that simple. Take notes, photos, and hikes. Watch your porch light at night (a fun way to confuse your neighbors is by staring at your porch light for an hour ion the evening). Who knows, one of us might make some new discoveries. Enjoy your wasps!

References

Alvarez, Juan Manuel (15 April 1997). "Chapter 26: Largest Parasitoid Brood". Book of Insect Records. University of Florida. Retrieved 3 September 2013.http://entnemdept.ufl.edu/walker/ufbir/chapters/chapter_26.shtml

Bin, Ferdinando, et al. “Tyloids in Pimpla turionellae (L.) are release structures of male antennal glands involved in courtship behaviour (Hymenoptera: Ichneumonidae).” International Journal of Insect Morphology and Embryology, vol. 28, no. 1-2, 1999, pp. 61–68., doi:10.1016/s0020-7322(99)00015-x.

Broad, Gavin. “Identification key to the subfamilies of Ichneumonidae (Hymenoptera).” 2015.http://nocturnalichs.myspecies.info/files/Ich_subfamily_key_April_2015.pdf

Burke, Gaelen R., and Michael R. Strand. “Polydnaviruses of Parasitic Wasps: Domestication of Viruses To Act as Gene Delivery Vectors.” Insects, vol. 3, no. 4, 2012, pp. 91–119., doi:10.3390/insects3010091.

Crankshaw, Owen S., and Robert W. Matthews. “Sexual behavior among parasitic Megarhyssa wasps (Hymenoptera: Ichneumonidae).” Behavioral Ecology and Sociobiology, vol. 9, no. 1, 1981, pp. 1–7., doi:10.1007/bf00299846.

Foster, M. A., and W. G. Ruesinki. “Influence of Flowering Weeds Associated with Reduced Tillage in Corn on a Black Cutworm (Lepidoptera: Noctuidae) Parasitoid, Meteorus rubens (Nees von Esenbeck).” Environmental Entomology, vol. 13, no. 3, Jan. 1984, pp. 664–668., doi:10.1093/ee/13.3.664.

Handwerk, Brian. “Oldest-Ever Bee Found in Amber.” National Geographic News, 2006.https://news.nationalgeographic.com/news/2006/10/061025-oldest-bee.html

Janzen, Daniel H. “The Peak in North American Ichneumonid Species Richness Lies Between 38 Degrees and 42 Degrees N.” Ecology, vol. 62, no. 3, 1981, pp. 532–537., doi:10.2307/1937717.

Khalaim, Andrey I. et al. “Mexican species of Exetastes (Hymenoptera: Ichneumonidae: Banchinae), with description of three new species.” Rev. Mex. Biodiv., vol. 83, no. 2, 2012.http://www.scielo.org.mx/scielo.php?script=sci_arttext&pid=S1870-34532012000200007

Maingay, Hilde M., et al. “Predatory and Parasitic Wasps (Hymenoptera) Feeding at Flowers of Sweet Fennel (Foeniculum vulgare Miller var. Dulce Battandier & Trabut, Apiaceae) and Spearmint (Mentha spicataL., Lamiaceae) in Massachusetts.” Biological Agriculture & Horticulture, vol. 7, no. 4, 1991, pp. 363–383., doi:10.1080/01448765.1991.9754566.

Mayhew, P. J. “The evolution of gregariousness in parasitoid wasps.” Proceedings of the Royal Society B: Biological Sciences, vol. 265, no. 1394, July 1998, pp. 383–389., doi:10.1098/rspb.1998.0306.

Nel, A. et al.. “An aulacid wasp in the Lowermost Eocene amber from the Paris Basin (Hymenoptera: Aulacidae).” Geologica Acta, vol. 2, no. 1, 2004, pp. 67–74.http://www.redalyc.org/pdf/505/50500110.pdf

Nuttall, M. J. “Pre-Emergence Fertilisation Of Megarhyssa Nortoni Nortoni (Hymenoptera : Ichneumonidae).” New Zealand Entomologist, vol. 5, no. 2, 1973, pp. 112–117., doi:10.1080/00779962.1973.9722978.

Owen, T. “Wasps of 2016.” Amateur Anthecologist, 2016.http://www.amateuranthecologist.com/2017/01/wasps-of-2016.html

Ridley, Mark. “Clutch Size and Mating Frequency in Parasitic Hymenoptera.” The American Naturalist, vol. 142, no. 5, 1993, pp. 893–910., doi:10.1086/285579.

Sandanayaka, W.r.m., et al. “The effect of mating behaviour on progeny sex ratio ofMastrus ridens(Hymenoptera: Ichneumonidae), a biological control agent of codling moth.” Biocontrol Science and Technology, vol. 21, no. 4, 2011, pp. 485–496., doi:10.1080/09583157.2011.552103.

Shcherbakov, Dmitry E. “Permian ancestors of Hymenoptera and Raphidioptera.” ZooKeys, vol. 358, 2013, pp. 45–67. doi: 10.3897/zookeys.358.6289

Stepahin, Chris. “Crystalline Tube Gall.” The Nature Niche, 17 Oct. 2014, thenatureniche.com/2014/10/17/crystalline-tube-gall/.

Stephen G. Compton, Alexander D. Ball, Margaret E. Collinson, Peta Hayes, Alexandr P. Rasnitsyn, Andrew J. Ross. Ancient fig wasps indicate at least 34 Myr of stasis in their mutualism with fig trees. Biology Letters, 2010; DOI: 10.1098/rsbl.2010.0389

Suzuki, M., et al. “The virus-like particles of a braconid endoparasitoid wasp, Meteorus pulchricornis, inhibit hemocyte spreading in its noctuid host, Pseudaletia separata.” Journal of Insect Physiology, vol. 54, no. 6, 2008, pp. 1015–1022., doi:10.1016/j.jinsphys.2008.03.013.

Tao, Li et al. “Effect of the trap color on the capture of ichneumonids wasps (Hymenoptera).” Rev. Colomb. Entomol., vol. 38, no. 2, 2012.

Ueno, Takatoshi. “Adaptiveness of sex ratio control by the pupal parasitoid Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size.” Evolutionary Ecology, vol. 12, no. 6, 1998, pp. 643–654., doi:10.1023/a:1006577314205.

Vinson, S. Bradleigh. “Courtship Behavior and Evidence for a Sex Pheromone in the ParasitoidCampoletis sonorensis(Hymenoptera: Ichneumonidae)1.” Environmental Entomology, vol. 1, no. 4, Jan. 1972, pp. 409–414., doi:10.1093/ee/1.4.409.

Useful Websites

Microgastrinae Wasps of the Worldwww.microgastrinae.myspecies.info/

Genera Ichneumonorum Nearcticae American Entomological Institutewww.amentinst.org/GIN/

Pteromalidae UC Riversidewww.faculty.ucr.edu/~legneref/identify/pteromal.htm

Universal Chalcidoidea Database. Natural History Museumwww.nhm.ac.uk/our-science/data/chalcidoids/torymidae.html

Special thanks to John Jacob @ Old Sol Bees for letting me use his microscopes.

Aculeate wasps of Southern Oregon pt 2

noreply@blogger.com (Travis Owen) — Sat, 04 Nov 2017 02:58:00 +0000

(This is Part 2 of Aculeate Wasps of Southern Oregon, see Part 1 by clicking here)

Cuckoo wasp, Chrysis sp.

Wasps are amazing. I love the wasps that everyone (in the colloquial sense) hates, yellowjackets, hornets, and paper wasps. Vespid wasps are probably the most evolutionarily advanced of the wasps, since in the Hymenoptera eusocialism appears to be the trend. The wasps that truly fascinate me are the hundreds of thousands of solitary predatory and parasitoid wasps that live their lives without much notice. The lives of the solitary wasps are so incredibly diverse there is no vertebrate comparison. What I present to you is but a small sampling of aculeate (stinging) wasps I have encountered in Southern Oregon. The term "stinging wasps" is conflicted because not all are capable of stinging people. Older lineages of wasps (the true parasitoids) did not possess stingers but rather had ovipositors. Ovipositors are elongated egg-laying apparati used to lay eggs on or within a host, such as a caterpillar. Aculeate wasps differ from their ancestral relatives, since their ovipositors have evolved into stingers. Venoms which initially were used to paralyze hosts now possess pain-inducing molecules, among other things. As stated previously in Part 1, most solitary wasps are reluctant to sting, and are practically harmless. Don't kill them.

Sphex lucae visiting wild carrot, Daucus carota

Sphecidae

The Sphecidae are solitary wasps which are easily differentiated from the Crabronidae by the elongated first abdominal segment, giving rise to the common name thread-waist wasps. Most of the wasps are predatory, while a small percentage are kleptoparasitoids which will take food from others for their young. These wasps may seem daunting to some, but they are hard pressed to sting. Even after handling several, even attempting to get stung (who does that?!), I was disappointed when they fled or just wouldn't sting me, those jerks. Being solitary wasps, their sting is better adapted to stinging soft bodied larvae or thin membranes between the plates of armored prey than stinging an annoying human.

Ammophila sp.

Ammophila adults are commonly seen visiting flowers where they are native. Females dig underground nests with a single cell chamber, occasionally two. They prey on moth caterpillars or caterpillar-like sawfly larvae, which are paralyzed by the sting of the female, and then dragged into the nest. Sometimes many small caterpillars are progressively delivered to the growing larvae, but more often a single large caterpillar is deposited with an egg and the nest is sealed for good until the new wasp emerges. One particular trait of these wasps is that they like to disguise their nests after they are sealed, or even before they bring prey or lay eggs. I once watched as a large female brought small pieces of bark, or large in relation to the wasp, to conceal a nest entrance. This behavior may discourage would-be predators or parasites from finding the precious larvae.

Ammophila sp.

Ammophila, like other predatory sphecids, have large powerful mandibles. Their heads may also be big, correlating to the size of their preferred prey, since the muscles that allow them to carry their prey are in their heads.

Ammophila sp. in the subalpine zone of Mt. McLoughlin

When I climbed up Mt. McLoughlin in August, I found Ammophila in the subalpine zone, just above the tree line. Ammophila prefer to nest in sandy soil, and that is what is found up there. Soils high on mountains are usually rudimentary, high in mineral content and low in organic matter, and perfect for many ground nesting wasps and bees (both of which were highly active during my hike on Mt. McLoughlin).

Sceliphron caementarium

Black and yellow mud daubers are just as familiar or more familiar than Ammophila, but are probably more recognizable for their nests than the wasps themselves. Mud daubers build mud nests on exterior surfaces, and they aren't picky about what kind of surface. However, they have an affinity for hidden and protected nesting places such as inside empty bee hives or under siding. They have even been known to attach nests to airplanes or shipping containers, and are currently found in every continent except Antarctica, including many remote islands. Nests are usually found close to sources of mud such as rivers or springs. This is in contrast to potter wasps (Eumeninae) which create their own mud by collecting water and mixing it with fine mineral medium to create their urn-shaped nests. Nests in different regions vary based on the locally available mineral composition of the available mud. Mud nests survive best in protected niches because rain can severely weaken the mud and put the developing wasps at risk, though from the prepupal stage the wasps surround themselves in a cellophane-like casing which is at least partially water resistant (personal observation).

Sceliphron caementarium

Mud dauber nests are formed from a grouping of one to several cell chambers, coated in totality in a protective layer of mud. Parasites and parasitoids infiltrate the mud nests despite the best efforts of the nest builder. Various fungal and (assumed) bacterial infections kill developing larvae and pupae, as well as death of larvae and pupae by kleptoparasitic or predatory interlopers (including other hymenopterans and vertebrates). Survival of developing mud daubers is hindered by these pests but countered by the many eggs laid per female each year, at least 25 cells in large nests (possibly multiple nests per female). I have often found that even in heavily infested nests, at least one or two larvae survive.

At the end of 2016 I collected several S. caementarium nests. Several had begun to hatch out, so I broke one apart to reveal the wasps in their pupal casings. In normal circumstances the wasps have to chew their way out of the cellophane-like casing and then proceed to chew their way through the wall of the mud nest to escape. In this situation I observed a young wasp that was very active within the casing, while the others were visibly inactive and presumably not mature enough to emerge yet. Perhaps it was the warmth of my hand, or maybe just by chance, but I was extremely fortunate to observe a mud dauber emerge for the first time. I was nearly overcome with emotion reminiscent of the birthing of my two daughters. A life had emerged into the world for the first time, and it moved me.

Contents of Sceliphron caementarium nest

Black and yellow mud daubers prey on spiders, usually orb weavers and crab spiders (varying regionally, I suppose). As many as fifteen spiders will be captured, paralyzed, and stuffed into each cell for the developing larvae to feast on. I dissected a nest and discovered the spiders pictured above, mostly crab spiders (Thomisidae). They were all presumably alive, and a few could move but only slightly, the venom apparently having either worn off or not being fully effective from the beginning. I discovered larvae in different stages of development, different instars. S. caementarium is multivotine, meaning they have multiple generations per year, and that nests are prepared all year long so long as the climatic conditions are favorable. I placed these larvae, and spiders, into a small closed wooden box (~1 square inch) and upon inspection a month later discovered webbing suggesting at least one of the spiders had regained enough mobility to spin silk. I wonder if, despite the shortcomings of the venom or maybe misplacement of the sting, spider prey is kept from harming the wasp larvae by the immobilizing affect of cell confinement.

Great golden digger wasp male, Sphex ichneumoneus, visiting narrowleaf milkweed, Asclepias fascicularis

The great golden digger wasp is a common sphecid native to nearly all of the lower 48 states, parts of Canada, and Mexico. As the name implies, the species is ground nesting. Up to six katydids are paralyzed and placed into underground cells. Birds, particularly house sparrows (Passer domesticus) have been observed (not by me) to startle digger wasps forcing them to drop their katydid prey, whence the birds claimed the katydids for themselves. A small percentage of females will forgo the creation of their own underground nest and proceed to provision another digger wasp nest, saving the labor of digging the nest herself. Females provision the same cells instead of just share the same nest entrance. Similar behavior in ancient wasps is theorized to have been a precursor to hymenopteran eusociality. If the joint nest mates should ever meet they will fight, but if they never encounter one another whoever lays the first egg and seals the nest will "win" the nest while the other is forced to dig her own nest or forfeit her reproductive success.

Sphex lucae visiting wild carrot, Daucus carota

Another species of Sphex (less studied) native to the Western states (From WA in the north, south to CA, east to TX). Sphex are known to prey only on katydids (Tettigoniidae), and each species of wasp preys on a single species of katydid. The wasps themselves have been known to be hosts to twisted-wing parasites (Strepsiptera). Strepsiptera are true parasites, wingless females live their entire lives embedded between abdominal tergites of many hymenopterans. Sphex are also victim to various nest parasites, yet some species are known to build false burrow entrances (accessory burrows) to mislead and distract interlopers from the true nest. Some crabronids (i.e. Bembix and Philanthus) are also known to dig accessory burrows. Ammophila, another ground nesting sphecid, misleads would-be nest invaders by depositing excavated soil away from the nest entrance.

Grass-carrying wasp female, Isodontia sp., visiting parsley flowers

Many wasps are fairly inconspicuous when creating nests. Not Isodontia, known as grass-carrying wasps, a tip of their preferred nesting material. Isodontia are cavity nesting wasps which use large chunks of grass to separate cells and cap nest entrances, and are often seen carrying pirces of grass larger than themselves. Their typical nesting sites are hollow stems, but they will also nest in other cavities in wood, or in some cases, umbrella poles (or other manufactured human things). Their prey are Orthoptera, typically Gryllidae or Tettigoniidae. One to twenty prey items are placed in each cell, varying between species of wasp and probably the size of the prey.

Prionyx sp. (atratus or subatratus) on rabbitbrush, Ericameria nauseosa, on Upper Table Rock

I discovered an individual Prionyx (either atratus or subatratus, the only all black Prionyx in the US) late in the day visiting a rabbitbrush plant at the summit of Upper Table Rock. The sun was setting and the wasp seemed to be wandering around on the shrub for no particular reason. Like some of the other sphecids I've showcased, Prionyx preys on Orthoptera, particularly short-horned grasshoppers (Acrididae), taking both older nymphs and adults. They usually capture and subdue prey, paralyzing it with its sting, then proceed to excavate a burrow after prey is acquired. A single cell is created at the end of each burrow. A single prey item is normally delivered per egg/cell, since chosen prey are typically large in relation to the wasp.

Pompilidae

Calopompilus pyrrhomelas visiting threadleaf milkweed, Asclepias fascicularis, at the base of Lower Table Rock

Some of my personal favorite solitary wasps are the spider wasps, or pompilids. I love all the wasps, but spider wasps are rock stars! Most pompilids hunt spiders, normally taking them head on, paralyze them, drag them into a hole, and seal them in with an egg. The living spider is eaten while the wasp sucks the life force right out of it. Another notable feature is that the sting of some pompilids is known to be very painful, the most painful hymenopteran sting award going to the large tarantula hawks (Pepsis spp.) of the American Southwest. Despite the pain, practically no lasting damage is done (except maybe for a damaged ego), as opposed to eusocial wasps and bees whose venom breaks down cell tissue and other unfortunate effects. I haven't been stung by any of the spider wasps, though it is truly on my wish list.

Calopompilus pyrrhomelas visiting threadleaf milkweed, Asclepias fascicularis

In Southern Oregon the largest pompilid I've encountered is Calopompilus pyrrhomelas, a smaller cousin of the tarantula hawks. They prey on a single spider species, as most spider wasps do. This spider wasp takes folding-door spiders, Antrodiaetus (probably A. pacificus), but is said to face them directly in the spiders own burrow. If the wasp is successful, the spiders burrow becomes its grave and the wasp adheres an egg directly between the spiders legs and seals the cell. The spider, doomed, and a new wasp lives. Calopompilus is seldom found on flowers, though they may have an affinity for milkweeds like their larger cousins in the South (some milkweeds may be primarily pollinated by large wasps, like pepsine wasps), and also for rabbitbrush (Ericameria nauseosa) as the host records demonstrate.

Auplopus sp.

Most pompilids nest in the ground, but a few nest in nests constructed above ground. Auplipus create mud nests, a jumble of mud cells, in protected locations such as under leaves or siding, or more often in preexisting cavities created by beetle larvae or other wasps. Some may even nest in abandoned mud dauber nests. Some species remove the legs of their spider hosts in order to feed on the hemolymph, or to make transport of the paralyzed spider easier. Eric Eaton has written a nice piece on these wasps, see it here.

Evagetes sp. in the subalpine zone on Mt. McLoughlin

It seems that the deeper one delves into the world of wasps, the more one must expect the unexpected. Many hymenopteran families have their very own black sheep, and Evagetes (Pompilinae) is one of the black sheep in the Pompilidae. They are social parasites, or kleptoparasites, who lay their eggs in the nests of other pompilids. Pompilids in the subfamily Ceropalinae are also known to be kleptoparasites, as well as paper wasp (Polistinae) mimics.

Evagetes sp.

Female Evagetes seek out completed pompilid nests, perhaps of a specific host species. I am not clear how this is done, perhaps by a scent or some other means. When a host nest is discovered, the female digs down into the chamber with the spider and host egg. Sometimes the egg is destroyed by the parasite, other times the host egg is eaten by the Evagates larvae which hatches prior.

Mutillidae

The mutillids are often confused for ants, since the females are hairy and wingless. The common names offer no help names such as "velvet ant" or "solitary ants". They can be differentiated from ants, which have one or two nodes between the mesosoma and metasoma, absent in wasps and bees. Another common name, "cow killer", is usually reserved for a single species, Dasymutilla occidentalis, which is known to have a very painful sting.

Dasymutilla sp. female

Female velvet ants are ectoparasitoids or kleptoparasites of a variety of other wasps, bees, and perhaps ants or their inquilines. Some mutillids are nocturnal, while the majority are probably diurnal. The combination of their painful sting, bright colors, heavily armored bodies, and an assortment of other aposematic defenses means they are usually left alone, even when boldly trekking across bare ground. Most are parasitoids of solitary unguarded larvae or their provisions, but some seek hosts within social nests of halictid bees or even honey bees in some European species. In the case of eusocial hosts, the wasps make their way through the bee guards, by just walking right in or killing them, and parasitizing the closed cells containing young pupae. An egg is laid, and the mutillid larvae feeds on the host pupae until pupating itself.

Mutillidae male

Male mutillids often look nothing like their female counterparts. They can be so different than females in appearance that some have even been wrongly classified as different species. Males also look suspiciously similar to wasps in the families Bradynobaenidae and Chyphotidae, though none are closely related.

Sapygidae

Eusapyga sp.

Sapygid wasps are Batesian mimics of eusocial Vespidae, though much less commonly encountered. They are ectoparasitoids or kleptoparasites of bees, mostly in the Megachilidae, though a few host records reveal eumenine wasps as hosts to some genera outside the US. Typically the newly hatched first instar sapygid larvae, which hatch prior to the host egg, puncture and consume the contents of the host egg (known as hospicidal behavior) before feeding on the provisions provided by the host (i.e. pollen).

Eusapyga sp.

Sapygids visit flowers, mostly Apiaceae and Asteraceae where nectar is often easily accessible. Like other parasites and predators of bees, flowers are often not far from nests of hosts. Megachilid bees, the most common hosts, also tend to visit flowers with easily accessible resources. I have never personally seen (or was aware of) sapygids visiting flowers. I had found one near a large apiary, a place I have observed several species of megachilid bees including Megachile and others.

Chrysididae

Chrysis sp.

The Chrysididae is a large and varied family with common names such as cuckoo or jewel wasps, for the life styles or physical appearances of some, respectively. They are unique among the Hymenoptera by possessing a specialized ovipositor which evolved solely within the Chrysididae. The ovipositor, much unlike the parasitic Ichneumonidae and related parasitoids, is made from the last several segments of the abdomen and is telescoping. Some chrysidids are kleptoparasitic, laying eggs in the nests of other hymenopterans to feed on their stored provisions (and killing the host larvae), or true parasitoids by feeding on the larvae or pupae of a host (i.e. pupating moths).

Chrysis sp.

Most chrysidids lack the ability to sting, and rely on their stout and heavily armored bodies as defenses against stinging hosts. Many of the larger species are metallic blue, green, or sometimes red, while smaller species are dark metallic green or black. They can sometimes superficially resemble torymid wasps (Torymidae), blow flies (Calliphoridae), or some bees (metallic green bees in the Halictidae and Megachilidae), but the short legs and antennae on chrysidids is somewhat distinguishable to the trained eye.

Chrysis sp. visiting Daucus carota

Chrysidid adults visit some flowers for nectar, most notably flowers in the carrot family (Apiaceae) which have many small flowers in clusters with easily accessible nectar.

Bethylidae

Epyrinae

Bethylids are very small wasps related to the chrysidids. They are ectoparasitoids of beetle or moth larvae. Females possess stings which they use to permanently or temporarily paralyze hosts so they can attach an egg to the hosts body. In the case of some larval hosts, female bethylids have been observed removing hairs from the hosts body in order to achieve good egg adhesion. Bethylid larvae sometimes live part of their lives as ectoparasitoids, but in later instars become endoparasitoids and finish off the host from within. Female bethylids sometimes feed on the hemolymph of the host, even killing them. This behavior may contribute to the fitness of the females, increasing their egg laying capacity. Depending on the species, females may lay between 50 and 100 eggs in their lifetime.

Epyrinae, with a US quarter for scale

I discovered the bethylid pictured above inside a highly active honey bee hive. The small wasp, nearly the size of the antennae of the much larger honey bees, was completely ignored until I caught sight of it. Some bethylids are known hosts of Cucujoidea or Pyralidae, the families containing hive pests small hive beetles (Aethina tumida) and greater wax moths (Galleria mellonella), respectively. Bethylids are also parasitoids of certain house pests, such as carpet beetles (Dermestidae).

Tiphiidae

Paratiphia sp. visiting Daucus carota

Tiphid wasps, sometimes known as flower wasps, are parasitoids of ground nesting beetles (mostly Cicindelinae and Scarabaeoidea) as well as some ground nesting bees and wasps. Host larvae are temporarily paralyzed, then eggs are attached to the underside of the host. Adult females sometimes feed on the host themselves in order to supplement their diet of flower nectar and honeydew.

Paratiphia sp. visiting Daucus carota

There are many more species of wasps in Southern Oregon, many that I haven't encountered myself. This is good, because it leaves the future to discover these treasures and refresh my always growing love and passion for wasps and other hymenopterans. Return soon and learn about the fascinating and more primitive parasitic wasps, such as the ichneumonids, diapriids, and pteromalids! Also to come, more bees! Stay tuned while I try to keep the approaching winter from being boring and drab.

Evagetes sp.

Aculeate Wasps of Southern Oregon pt. 1

noreply@blogger.com (Travis Owen) — Sun, 15 Oct 2017 16:00:00 +0000

Stelionia male visiting Monardella sp. for nectar

Wasps have a bad reputation. Ask ten people what they think of wasps and seven of them will tell you they do not like wasps. This is what I learned after I created a survey earlier in 2017 (simply entitled Wasps. You can still participate in the survey here). The survey was distributed via social media, email, and word of mouth. The majority of the participants considered themselves to be nature enthusiasts and gardeners, though academics and beekeepers also made up a large percentage, and lastly a small percentage considered themselves none of these things. Although the survey had some issues, and lacked some of the qualities of a professional scientific study, what was clear was that wasps as a whole are not viewed in the best light. This was reinforced in the end of the survey, when I allowed participants to write a comment relating to wasps. Some of the comments showcased knowledgeable participants who noted the beneficial aspects of wasps, including the non-stinging parasitoids (not featured in this article), and the non-aggressive nature of many of the solitary wasps. Most of the responses, however, were less pleasant with comments like I like bees but not wasps or yellowjackets are a**holes. My mission is to confront the prejudice and lay out the reality of wasps which is both fascinating and occasionally obscure (and oh so alluring to me). There is much to learn of wasps.

Spider wasp, Evagetes sp. (Pompilidae) with relatively long sting. Despite the intimidating appearance of the sting, these wasps are far more likely to flee than try to sting, even when I captured one with my hand. Evagetes are kleptoparasitoids of other pompilids, and do not build their own nests.

Being stung is likely the primary concern most people have regarding wasps. When it happens, it can be painful and unpleasant. Stings from bees and wasps can instigate allergic reactions ranging from itching and swelling, hives, or on rare occasions, life threatening asphyxiation. The CDC attributes around 100 deaths per year in the US to stings by insects and other arthropods (i.e. scorpions), though other sources estimate 50 deaths a year, which is one out of every six million people in the US (or one out of every three million for all stinging deaths). This doesn't discern whether the deaths were a result of allergic reaction. One common misconception is that for those who are allergic to honey bee stings they are also allergic to stings from wasps. Venom chemistry is complicated and differs significantly between wasps and bees, as well as many considerable differences between the venoms between different species of wasps, bees, and ants. No two are exactly alike, and most are dramatically different. That said, being allergic to both honey bee and paper wasp stings, for example, is extremely unlikely. For more on the science of stinging Hymenoptera and venom, I encourage the reader to see the work of entomologist and researcher Justin Schmidt.

Eusocial Vespidae

Am I ready for my closeup? The face of a bald-faced hornet, Dolichovespula maculata

The most notorious, or rather infamous, of all wasps are the wasps in the family Vespidae. This family includes eusocial wasps such as yellowjackets (Vespula), aerial yellowjackets (Dolichovespula), true hornets (Vespa), and paper wasps (Polistes and Mischocyttarus), as well as an array of non-aggressive solitary wasps. These wasps interact with people in a variety of ways, nesting under eves in the case of the paper wasps, nesting in lawns in the case of yellowjackets, or in all cases crashing your picnic or BBQ party. Social vespids are all predatory. Usual prey include flies, caterpillars, and many garden pest species, but they may also hunt larger arthropods or be attracted to carrion. Unlike the solitary vespids, chewed up pieces of the prey are brought back to the nest rather than a complete paralyzed yet living prey item.

Yellowjacket sting, Vespula pensylvanica, under magnification

Eusocial bees and wasps inflict not only painful stings, but inflict tissue damage as well. Solitary wasps and bees, by contrast, are often reluctant to sting but when they do it is often weak. But the majority of solitary bees and wasps that are willing to sting, regardless of causing any notable pain or not, do not cause any lasting damage. The significance behind this is that eusocial wasps and bees must protect their nests which are contain their kin, not to mention any food stores they have acquired. Solitary bees and wasps, in contrast, have much less invested into their nests and can usually relocate and build a new nest.

Vespula pensylvanica, up close and personal

The most hated wasps, it seems, are the yellowjackets. True yellowjackets, Vespula spp., are eusocial ground nesting wasps native to every state of the US and much of Canada. They are easily disturbed and meet any perceived threat with aggression. Their habit of nesting in lawns has brought them too close to home, literally, and are often cause for concern for those with small children or dogs (as a father of two young daughters, I don't blame them! I don't usually condone the killing of wasps or bees, but if one should need to get rid of a nest then wait til the cover of darkness and pour a bucket of soapy water into the nest entrance.) Yellowjackets, like honey bees, live in nests with workers and a single laying queen (excessively large nests of yellowjackets in the southern states may have two or more queens). Queens produce pheromones, much like honey bee queens, which alter worker behavior by suppressing the instinct of workers, all female, from laying eggs. Pheromones also prevent other queens from taking over since they are just ever so slightly specific to each individual queen. Nests are composed of paper, and are structured into several layers of downward facing hexagonal cells in horizontal combs. They hunt all types of arthropods and are also attracted to carrion (or hamburgers). Prey is not stung, but rather bitten apart. What might appear to be stinging is actually just respiration, and the use of the abdomen as an extra limb used to grapple with prey.

European paper wasp, Polistes dominula

Paper wasps are similarly notorious and often interact with humans due to their nesting preferences. Paper wasps nest in elevated and sheltered locations, such as within tree hollows, or more familiarly, under eaves. The nests are made from paper, like all domestic social vespids, but unlike Vespa, Vespula, and Dolichovespula the comb is exposed and not enveloped in a paper shell. Nests are also relatively small, rarely growing larger than eight inches across. In my experience, paper wasps in Oregon are very docile and do not attack except under extreme disturbance (like the time I decided to pry a nest off with a stick). Otherwise, they seem to flee or just fly around rather than stinging. I also find the sting, when it does happen, to be mild compared to honey bees or yellowjackets.

Green eyed Polistes dominula male nectaring on puncture weed, Tribulus terrestris

Polistes dominula is a European introduction, now widespread in the West and elsewhere in the country. They are a highly adaptable species and less particular about nesting sites than native paper wasps. There is reason to believe they displace native species by dominating the most favorable nest sites.

Polistes dominula wing and abdomen under a microscope

Polistes dominula, and all vespid wasps, have pleated wings. The wings appear thin precisely because the forewings are folded in half when not in flight! This is thought to prevent damaging the fragile wings while in close quarters with nest mates. This trait is also seen in a few solitary bees, flies, and a few other flying insects (all wasp mimics). Most Hymenoptera also have fine hairs covering the wings which may serve multiple purposes including water repellency, reducing drag, thermoregulation, and perhaps other unknown functions. The length and quantity of hairs varies between different wasps and bees, perhaps catering to their varying lifestyles.

Honey-like substance in Polistes dominula nest

Paper wasps, and perhaps other eusocial vespids, visit flowers for nectar which is a source of carbohydrates (necessary for an active lifestyle if you are an adult wasp or bee). Occasionally this resource is brought back to the nest and stored for later use. It was very thick, almost gelatinous and not nearly as viscous as true honey, but was very sweet. A distant relative of the paper wasps, Mexican honey wasps (Brachygastra mellifica) are vespid wasps which build large hanging paper nests in Southern Texas and Mexico. They collect large quantities of nectar and process it into large quantities of a honey like substance. Occasionally the wasp honey is collected, but unfortunately its collection requires the complete destruction of the nest. (By the way, if you happen to have any I'd love to taste it!)

Polistes aurifer, the golden paper wasp, is native to the Western US and Canada

Some paper wasps use a form of facial recognition to communicate with nest mates and understand hierarchy. Species with less detailed facial patterns do not communicate this way, and instead have comparatively larger olfactory sensors. Polistes dominula, P. fuscatus, and I suspect P. aurifer, are some that have demonstrated facial recognition of nest mates. There is also some speculation that they can recognize the faces of people, though I could not find any literature substantiating that.

Polistes aurifer (above) and Apis mellifera (below)

Unlike yellowjackets and hornets, paper wasps are not aggressive towards honey bees. This is a concern for some beekeepers I have spoken to who attempt to justify their general prejudice towards wasps as a whole. In general, keeping honey bee colonies strong by the proper care and attention (i.e. IPM, feeding as needed, good genetics, etc.) will keep any wasp threat from subjugating the hive. Weak colonies are not well equipped to defend their home, thus thieves or predators may freely enter. Paper wasps, unlikely to prey on honey bees themselves, will rob honey from weak or dead colonies. I would argue that the wasps are merely a symptom, but not the cause, and actions should be taken to build up the strength of the hive rather than eliminating the perceived wasp threat.

Mischocyttarus flavitarsis, Western paper wasp

The largest paper wasps in Oregon are in the genus Mischocyttarus. They are differentiated from Polistes by having a thin elongated waist and reduced front legs, often standing on the center and rear pairs of legs. Nests are similar in appearance to those of Polistes, and my only observation is that they are less abundant than Polistes, while P. dominula is the most abundant. The long legs of all the paper wasps I have observed allow them to land directly on the surface of still water by spreading their legs and utilizing the surface tension.

Polistes dominula female, magnification showing the waist (petiole)

The narrowed but not elongated waist of Polistes wasps, as seen in the photo above, allow lay observers to differentiate them from the related Mischocyttarus. Relying on markings and patterns alone is not a reliable method of identification. For example, Polistes aurifer and Mischocyttarus flavitarsis females bear nearly identical markings and patterns in Southern Oregon, though perhaps one or both bear more distinct markings in other regions where different phenotypes take up residence.

Solitary Vespidae

Small mason wasp, possibly an Ancistrocerus spilopterus female (Eumeninae)

Solitary vespids are often very similar in appearance to their eusocial and semi-social (and usually more aggressive) cousins. The Vespidae contains five subfamilies, two are eusocial or semi-social (Vespinae and Polistinae, respectively) while the remaining are solitary. So far only known in California, the Euparagiinae is rare. I find the Masarinae to be particularly interesting as solitary females provision mud nests with pollen rather than insects. Maserine wasps collect pollen in their crop and regurgitate it into the nest, similar to masked bees (Hylaeus spp., Colletidae). The genus Pseudomasaris (Masarinae) is known to occur in Southwestern Oregon, though I have not seen one yet (to my knowledge). I'm keeping my eyes out, they are active in early summer and superficially resemble yellowjackets.

Honey bee, left, and mason wasp, right for scale

The Eumeninae is the most common of the solitary Vespidae. In general they are smaller than their eusocial cousins, but many bear a superficial resemblance with yellow and black patterns. Others have patterns that are black with either red, white, or orange. Despite their similarity to their occasionally aggressive extended family, the solitary vespids are quite reluctant to sting and will swiftly flee if a threat is perceived. Those who have managed to somehow get a sting from one of these wasps report disappointment in such a weak defense. Their first line of defense is aposematic, the bright showy colors resemble yellowjackets or other vespids with painful stings.

Eumenine wasps build nests out of mud in cavities, or as freestanding mud pots attached to something off the ground (i.e. house siding, grass, rocks). Many of the eumenines build nests in preexisting cavities such as deserted tunnels in wood or in the case of the above photo in old honey wax comb. Unlike many solitary bees which build linear nests in tunnels, mason wasps are more flexible in the nest structure. Mason wasps cap cells with mud. Potter wasps, however, build external structures that resembles pots with fared constricted entrances. In either case, the cells are stuffed with one large or many small paralyzed caterpillars and an egg, then left to their own devices.

Ancistrocerus sp. male

In 2016 I collected some mason wasp nests built in old honey bee comb to rear them and attempt to identify what emerged. Wasps begun to emerge as early as February (2017), starting with male Ancistrocerus as seen on my finger in the above image. Eumenine wasp eggs hatch shortly after they are deposited, within days to weeks, and quickly devour their paralyzed caterpillars. They enter winter as prepupae and remain that way until pupating the following year. I suspect temperature plays a large role in their pupation and emergence. At the time of the emergence of those first few Ancistrocerus males, there was still some remnant snow on the ground in the Rogue Valley. Secondly, the screened jar which held the nest was in my truck where it surely experience higher mean temperatures than outside, not to mention much drier conditions.

Showy milkweed, Asclepias speciosa, with a mason wasp

Mason and potter wasps are common flower visitors. Although they don't provision nests with nectar or pollen like some of the other vespids, adults visit flowers for nectar as it is a readily available source of carbohydrates. Flowers are also occasionally the site of feeding caterpillars or other favorable larvae, so may also be where adult eumenine females might search for prey.

Ancistrocerus tuberculocephalus male

Male eumenines do not provision nests, and do not hunt caterpillars. They do visit flowers for nectar, I captured a Ancistrocerus tuberculocephalus male foraging on deerbrush, Ceanothus cunneatus, in late spring. The best thing about the males is that they help create more females! Like all aculeate hymenopterans they can be identified by the extra abdominal segments which enable them to bend their abdomen to mate with females, and for an extra segment in their antennae, often curled at the tip.

Ancistrocerus sp., a mason wasp

Many observers have noticed that eumenine wasps have different types of nests, within cavities or as external structures. Common names serve to delineate them by nest type, yet, there is no well known taxonomic distinction between mason and potter wasps that I am aware of. I have observed them to be morphologically distinct, however, as they can be distinguished by differences of the petiole. The mason wasps that I am acquainted with have short petioles (see photo above) while potter wasps have an elongated petiole (see photo below) much like Mischocyttarus. are in general slightly larger than the mason wasps.

Eumenes male, a potter wasp

Perhaps the common names mason and potter wasp are not scientifically significant, though they do serve to describe the Southern Oregon species fairly. In my observations various eumenines are active from spring to fall, suggesting that many may be multivoltine, that is, having multiple generations throughout the year so long as good conditions prevail.

Eumenes verticalis

The Eumeninae are mostly predatory though a few Eurasian species are phytophagous (feed on plant tissue). Adults usually feed on nectar, or occasionally on hemolymph. Prey is captured for larvae, and is completely or partially paralyzed before sealing in a cell for the wasp larvae to feed on while developing. Species can be generalists when it comes to prey, and take caterpillars or beetle larvae from many families, while others can be more picky to the point of only taking prey of a single genus of moth or leaf beetle larvae, for example. Number of prey can also be variable for some species, such as one large prey item or several smaller prey items per cell. This variability is also found to be common in the Sphecidae and Crabronidae.

Crabronidae

Stelionia sp. female visiting Echinacea purpurea

The Crabronidae is thought to be closely related to the bees, the latter having evolved from the former. Ironically, some crabronids prey primarily on bees and other hymenopterans. Many crabronids are even mistaken for bees as they share some superficial traits. Having been fooled myself, I am certain of this. The Crabronidae prey on a wide variety of other insects and spiders. Prey are usually paralyzed and brought back to the nest for larvae to feed on.

Bembix americana female exiting her nest

Sand wasps in the genus Bembix are widespread in Oregon. They are called sand wasps because they nest directly in loose sand. I observed them on a sandy bank alongside Evans Creek (which meets the Rogue River) where a layer of hard packed silty soil formed a layer a few inches below the sand. The nest entrance itself fills with sand when the wasp isn't actively entering or exiting, so often appears as a small divot in the sand. Sand wasps are great diggers, diving into the sand with great ease, casting particles of sand behind them as one might imagine a digging dog.

Bembix americana hovering above her nest entrance

Bembix prey on flies, and don't seem to be specific about which types. Flies are paralyzed with venom and brought into the nest. Bembix provision cells progressively as their larvae develop, delivering up to twenty flies for each larva. Bristly hairs on their legs allow females to dig through sand with the two front legs while carrying prey with the remaining legs.

Steniolia sp. female visiting Echinacea purpurea

Steniolia dig nests in soil rather than sand, but are otherwise very similar to the sand wasps. I have also observed them to be slightly larger than Bembix, at least the species I've encountered. Nests descend six to seven inches ending in a single cell which, like Bembix, is provisioned progressively with flies as the larvae develops. Both genders of the genus cluster at night to share warmth, which is also when mating occurs (I haven't observed this, but I'd love to some day).

Steniolia male visiting Monardella sp.

Steniolia have very long tongues (probosces) when compared to other wasps, longer than many bees too. They have a particular interest in violet colored flowers such as thistles, some monardellas, and purple coneflower. Alternatively they are also known to be drawn to orange, yellow, or occasionally (but far less frequently) white flowers. Males can't sting, but are equipped with a double pronged pseudosting at the tip of the abdomen. Crabronids in general are quick and will readily flee if danger is perceived, but I hypothesize that if I were to capture a male with my hand he would poke me quite hard, perhaps hard enough to make me release him as a reaction. I have experienced a similar reactionary response when handling male wasps of all kinds who simply motion as if to sting, sans the pseudosting. Fear of being stung is powerfully integrated into the human psyche.

Philanthus gibbosus

Beewolves, Philanthus spp., are one of my favorite genera of all wasps. As much as I love bees, this bee-predator has charmed me. Beewolves are a group of ground nesting wasps which, unlike Bembix and Steniolia, mass provision cells before laying an egg and sealing the cell. Their primary prey are small bees, mostly but not strictly halictid bees (possibly a result of the abundance of halictid bees, some of which are semi-social), however beewolves are known to prey on other wasps and even occasionally other types of insects as well. Up to two dozen prey items will be packed into each cell before an egg is laid.

Philanthus gibbosus

I found Philanthus gibbosus nesting in near vertical banks along Evans Creek near the Rogue River. Sand wasps (Bembix americana) and osmiine bees were also nesting in close proximity. Some species of Philanthus are known to share a nest entrance but continue to provision their own nests within. Philanthus females are also known to line their cells with microbes which protect the developing larvae from potential pathogens. I wonder if this behavior occurs in other types of wasps or bees, yet awaits discovery.

Ectemnius sp. visiting Daucus carota

Ectemnius wasps prey on flies, mostly hover flies (Syrphidae). Unlike the aforementioned crabronids, Ectemnius nest in dried twigs or stems with soft pithy centers, or in holes in dead or decaying (soft) wood, varying between species. Cells are made from chewed pith and stuffed with flies, or maybe few to single large flies, depending on the species.

Ectemnius sp. in the subalpine zone on Mt. McLoughlin

In August of 2017 I hiked to the alpine zone of Mt McLoughlin halfway between Klamath Falls and Medford. I was fortunate to photograph Ectemnius nectaring on an unknown carrot relative (Apiaceae). Many other hymenopterans were active, from small halictid bees to sphecid wasps, pompilids, and chrysidids. Numerous dipterans (flies) were also active. Alpine and subalpine environments exhibit more extremes than lower elevations (i.e. climate), and life up there has evolved to survive the unique challenges presented. Plant life in the alpine and subalpine zones is particularly interesting, though notably more sparsely dispersed than the lowlands. This may lead one to think that life is less diverse in the mountain ecosystem, but that is a fallacy. Pollinators are quite numerous in the alpine zone, though not the same mix of pollinators found in the lowlands. Flies are very important pollinators of alpine and tundra environments since they are better able to cope with poor weather conditions, though many types of bees and wasps also flourish when the weather is favorable.

Tachytes sp. visiting narrowleaf milkweed, Asclepias fascicularis

I had originally mistaken this square-headed wasp for a honey bee, because honey bees were foraging on the milkweed in great numbers and the wasp was of a similar size and shape. Tachytes nest in the ground and provision each cell with Orthoptera adults or nymphs, primarily grasshoppers or katydids. Prey selection is somewhat specific depending on the Tachytes species. Some will provision every cell with a prey item before laying any eggs, temporarily storing the orthopterans in a special chamber off to the side while chambers are being excavated.

Tachytes sp., chilled not killed

Like many crabronids, this Tachytes was difficult to observe. To get a better look, and have a good chance of having it identified, I captured this wasp with an insect net and placed it in a jar. I placed the wasp in the refrigerator for ten minutes. The wasp was then chilled and temporarily immobilized, allowing me to get a good look. I brought it outside into the sun so it could warm up, after which it promptly flew off. This is a good technique for viewing many hymenopterans up close, though make sure to release the specimen as close as possible to where it was captured so it may carry on.

Aphid wasps, Stigmus sp.

Aphid wasps (Pemphredoninae) earn their name for their choice of host. Nests are constructed within existing holes in wood and mass provisioned with aphids. Don't confuse them with the other aphid wasps, Aphidiinae (Braconidae), which parasitize the aphids by laying an egg inside them.

Aphid wasps, Stigmus sp.

Initially I had confused these aphid wasps for masked bees, Hylaeus spp. (Colletidae), since they both lack pollen collecting hairs and are similarly small, less than 6mm in length. They are also both common visitors to wild carrot flowers, Daucus carota, an invasive weed here in Southern Oregon.

This concludes Part 1, follow the link to continue to Aculeate Wasps of Southern Oregon Part 2 which will include photos and bios of many lesser known aculeate wasps!

Apidae in Southern Oregon

noreply@blogger.com (Travis Owen) — Fri, 06 Oct 2017 05:39:00 +0000

Honey bees and a cuckoo bee, cousins in the family Apidae

The family Apidae contains the most well known bees, the honey bees, as well as bumblebees, carpenter bees, and a variety of other types of bees with interesting and diverse lifestyles. There are seven families of bees in the world (six found in the United States, the seventh is endemic to Australia), and of these, the Apidae is the most familiar. The Apidae vary greatly in size, appearance, biology, and lifestyle. The scope of this article is to introduce the reader to a few of the bees in this family found in Southern Oregon. Though the species highlighted here were all found in Oregon, they are representative of apid bees found throughout the country.

Golden paper wasp, Polistes aurifer, and a honey bee

Bees and wasps share a common ancestor. Bees diverged from wasps at least 130 million years ago, probably sometime in the Cretaceous. Wasps at that time were predatory, and were likely found around flowers to hunt for prey. Flowers were already producing protein rich pollen for other pollinators such as flies and beetles, with nectar-like secretions appearing in the Late Cretaceous. It is theorized that in some cases wasps started feeding on pollen as well as their insect prey, and the diets of some evolved to depend solely on pollen. Wasps, like modern wasps, have hair on their bodies, but bees developed branched hairs which are more efficient at holding pollen. This is the main feature that separates wasps from bees. Diet alone is not a reliable method of separating wasps from bees. South American vulture bees (Trigona crassipes, T. necrophaga, and T. hypogea) are members of the Apidae, and feed on carrion rather than pollen. Conversely, pollen wasps (Masarinae) are vespid wasps which provision nests with pollen rather than insects for their young. The eusocial members of the Apidae represent the most highly evolved of all bees and perhaps all wasps as well.

A little Bombus vosnasenskii worker steals honey from a honey bee hive

The genus Apis contains seven recognized species, including the giant honey bee (A. dorsata), the Asian honey bee (A. cerana), and the Western honey bee (A. mellifera). The Western honey bee has around 25 recognized subspecies, often referred to as races among beekeepers. In the US, only a handful of races are introduced, such as the Italian (A. m. ligustica), Carniolan (A. m. carnica), and Caucasian (A. m. caucasia) to name a few. The African subspecies, A. m. scutellata, has hybridized with many of the naturalized Eurasian subspecies, creating the Africanized bees which are restricted mostly to the southern states, limited by cold weather. Africanized bees have traits which help it counter one of the most important of honey bee pests, the varroa mite (Varroa destructor), and may help ensure the economic success of beekeepers in the South who suffer large winter losses from mite infestations.

Honey bee proboscis

Honey bees, as well as all other bees and wasps, have mandibles used for chewing and various other species specific tasks, and a proboscis. The proboscis, the bee equivalent of a tongue, is an assemblage of parts which form the sucking apparatus used to gather nectar or other fluids. The longest part of the proboscis is called the glossa, and it often has specialized hairs at the end. Some bees, such as certain osmiine bees (Megachilidae) have special curved hairs on the glossa used to collect pollen on certain host plants such as those in the Boraginaceae with concealed anthers.

Honey bee corbiculae

Honey bees, bumble bees, orchid bees, and stingless bees have corbiculae, or pollen baskets, which are specialized pollen collecting structures on the tibia of the hind legs of females. The outer hairs are long and curved inwards, while short soft hairs fill the interior. Pollen is mixed with bee saliva or nectar then packed tightly into the corbiculae. This is in contrast to most other pollen-collecting bees which collect pollen in a dry state in long dense straight hair clusters called scopae.

Honey bee sting

The honey bee sting, like the stings of all stinging wasps and bees, evolved from an ovipositor way back in the evolutionary timeline. Ancient female wasps (and many modern parasitoid wasps) laid eggs through the ovipositor. The rise of eusociality is thought to have correlated with the evolution of the sting, and venom, as a means of defense. The possession of a sting and venom may also have enabled wasps and bees to gather resources or hunt for prey out in the open, whereas many parasitoid wasps are relatively obscure and not often seen. The bright colors of many stinging wasps and bees is simultaneously an aposematic defense, a form of Müllerian mimicry (where distantly related species evolve to resemble each other), and in some cases, an example of Batesian mimicry (where generally harmless species resemble other not-so-harmless species).

Beekeepers and many others know that when one is stung by a honey bee, the bee dies. This is because the sting, which is barbed, remains in the victim with a pulsing sac of venom, as well as part of the bees digestive tract. The venom sac continues to pump the entire payload of venom if not promptly removed, best done by scraping. The bee dies, but this is usually of little consequence as the martyr is not a reproductive, that is, does not pass her genes, and is of greater use protecting the colony. This is altruism in the natural world. The vast majority of bees and wasps, however, do not have barbed stings and do not die after stinging because they retain their sting. The genus Apis, as well as a few South American eusocial vespid wasps (and a handful of stinging ant species), are the only hymenopterans with barbed stings. The myth that bees die after stinging and wasps can sting you til the cows come home turns out to be a fallacy.

Mated queen marked green

Honey bee colonies are composed of thousands of worker females and a single laying queen. The queen is larger than the others, and it is her sole purpose to lay eggs. When a new queen is hatched, she takes one to several mating flights where she mates with a few to many drones (males) in mysterious drone congregation areas, 50 to 100 feet above the ground. Once the queen has acquired enough semen, she never leaves the hive again and lays eggs until she is either killed, runs out of semen, or the colony swarms. As for the latter, the entire colony, a superorganism, reproduced through the act of swarming. This occurs usually during a time when resources are at their peak (i.e. spring) when the queen and loosely half the workers depart the current hive in search of a new place to call home. The swarm normally clusters on a tree branch, fencing, or any number of structures until scout bees find a suitable cavity or other protected location to build comb. The bees remaining in the original hive will raise a new queen from one of the last eggs laid by the former queen.

Apis mellifera caucasia male (drone)

Although the title of honey bee queen is purely an anthropomorphism, the queen doesn't actually rule anything except for the release of the semen within her ovaries. When an egg is fertilized, it produces a diploid female honey bee. Yet, the queen has the ability to lay an unfertilized bee which will develop into a haploid male, or drone. This trait is known as Haplodiploidy, and is consistent among most of the Hymenoptera (the taxonomic order which encompasses all bees, wasps, ants, sawflies, and horntails). I say most because there are some odd wasps which can produce females from unfertilized eggs. Male hymenopterans, having never possessed ovipositors since only females lay eggs, never developed stings. Honey bee drones, however, die after mating because their male organs are ripped from their bodies after intercourse with new queens. They also become paralyzed, and simply drop from the sky, their penis torn off. My problems aren't so bad, it seems. Most male wasps and bees, much unlike the honey bee drone, can mate repeatedly, and it is often the female which mates with a single male.

Eristalis tenax, drone fly (June)

Mimicry in the insect kingdom is widespread, particularly among flies. The drone fly, Eristalis tenax, is a mimic of honey bees. It is so named due to its strong resemblance to the honey bee drone. Evolution must know something I do not when it makes one harmless and edible creature mimic another equally harmless and edible creature. Flies, unlike bees and wasps, have only one pair of wings (bees and wasps have two pairs) and a pair of halteres behind the wings. Halteres are small knobbed appendages which allow flies to balance during flight.

Bombus vosnasenskii, the yellow-faced bumble bee

After honey bees, bumble bees (Bombus spp.) are the second most familiar bees. There are 46 species of bumble bees in the United States and Canada, over 200 worldwide. Around two dozen species have been found in Oregon, of which I have observed around five or six. They are quite different from honey bees, though there are also a few similarities.

Bombus vosnasenskii visiting Monardella sp., or coyote mint

Bumble bees are eusocial meaning they form colonies with distinct castes (reproductive and worker castes). Unlike honey bees, a bumble bee colony only lasts for a single growing season. While a honey bee queen can live up to around five years under ideal conditions, bumble bee queens live for one year. At the end of the colonies life cycle, new queens and males are produced and leave the colony to mate. The old queen and the workers of that season all die before winter sets in, while newly mated queens are the only ones to live til the following year, enduring winter sheltered in a protected location.

Bombus huntii

Bumble bees, as well as a handful of other large bees, are able to generate warmth by vibrating their flight muscles. For this reason new queens are able to forage for sustenance very early in the year, often late winter in my region when the earliest flowers are just beginning to bloom (winter heath, Erica carnea, is typically the only thing in bloom in late winter). Newly emerged queens seek out a nesting location which is usually underground (i.e. abandoned rodent tunnels) or within the thick tussocks of bunch grasses. Initially the queen will build a single wax urn and lay one to a few eggs inside. She will forage for nectar and pollen to nourish this first brood. When these first workers emerge as adults they will aid the queen and eventually take over foraging duties while the queen shifts gears to egg laying.

Bombus sp. forcing her way into a closed California poppy, Eschscholzia californica

Bumble bees are important pollinators with unique talents not exhibited by honey bees. First off, many species are larger and stronger than honey bee workers, sometimes with much longer tongues, and thus able to gain access to resources in flowers that are unavailable to honey bees. Additionally, the same mechanism that allows bumbles to warm themselves in cold weather allows them to gather pollen in some plants that is not able to be acquired by honey bees or other small bees. Sonication, colloquially known as buzz pollination, is when a bumble bee (or other bees with this ability) grip onto a flower or the anthers and vibrate their flight muscles to release the pollen. Many plants, particularly those in the nightshade family (Solanaceae) and heath family (Ericaceae), to name a few of economic importance (i.e. tomatoes, blueberries), require sonication for pollination as the pollen will not release from the anthers without it.

Bombus sp. visiting Heuchera sp.

Bumble bees are also very different from honey bees in that they do not communicate the locations of resources of resources as honey bees do (i.e. the waggle dance, a tool honey bees use to communicate the location of favorable flower patches relative to the sun). This lack of communication in bumble bee nests is useful for the use of bumble bees in agricultural settings, particularly greenhouses. While honey bees will forsake a target crop if a more preferred nectar/pollen resource is discovered beyond the farm, bumble bee foragers are all on their own to discover resources. Honey bees may ignore a target crop altogether by means of their communication with other foragers, but bumbles will not. This makes them ideal for greenhouse pollination efforts.

Bombus vosnesenskii on Hypericum perforatum

Bumble bees are adventurers. In my observations they have the most diverse collective palate of any native pollinator. In six years I have not witnessed a single pollinator visiting Klamath weed, Hypericum perforatum, which is a nonnative species from Europe, except for the bee in the above photo. This is just a single example of their dynamic ability of at least some species to adapt to changing conditions.

Bombus vandykei male visiting Echinacea purpurea

Most bumble bees are easily identified by observing which colors are on each tergite, the segments on the top side of the abdomen. In general they are easier for beginners to identify than other native bees due to their relative abundance and larger size. However, a few factors can complicate identification. Natural variability within the same species can occasionally give the impression to the observer that there are more species than the reality. Queens, workers, and males often have different patterns as well. For example, Bombus vandykei males can sometimes be completely yellow while queens and workers have a metasoma that is nearly all black except for a singe yellow band. B. vandykei and B. vosnasenskii females differ solely by the position of a single yellow band being one terga higher in the case of the former, an example of the importance of accurate observations. For most bees, a close look via good photos or an insect net and a hand lens are necessary for accurate identification. Some species are beyond the ability of the enthusiast, and require an expert with a microscope.

Robber fly, Laphria sp.

There are several bumble bee mimics in the region, mostly flies. Robber flies (Asilidae) and tachinid flies (Tachinidae) are the most common of the fly mimics. Not all of them are mimics of bumble bees, but several species bear a strong resemblance. Laphria spp., as well as other robber flies, are predatory and hunt other flying insects, including bees and other robber flies. Tachinid flies are parasitoids, their larvae feed on insect hosts and hosts in other orders including some arachnids (though hosts are species specific).

Xylocopa sp. female, a large carpenter bee, visiting hairy vetch, Vicia villosa

Often misidentified by lay observers for bumble bees, carpenter bees are common and conspicuous. They are the largest bees in Oregon, though unlike bumble bees they are solitary or occasionally primitively eusocial. Not only are large carpenter bees the largest bees in Oregon and much of the United States, but they also have some of the largest eggs of all insects, measuring over half an inch in length, though egg size is probably variable between species.

Xylocopa tabaniformis visiting clary sage, Salvia sclarea

Carpenter bees are similar to bumble bees superficially but several key features can help to tell them apart. The first, perhaps most obvious trait, is that the metasoma of Xylocopa is relatively shiny since the hair is considerable less dense than on bumblebees. Carpenter bees also lack corbiculae, or pollen baskets, and instead possess scopae, dense patches of elongated hairs on the tibia of the rear legs. Though I have observed that when visiting certain plants in the Lamiaceae (mint family) the pollen is deposited on the top of the mesosoma. I suspect that pollen is groomed into the scopae from other parts of the body though I have not observed this directly.

Male Xylocopa perched on a columbine

True to name, carpenter bees nest in wood. Unlike many other bees which nest in premade holes in wood created by beetle larvae or other insects, carpenter bees use their strong mandibles to create cylindrical nests in dead or living wood, occasionally bamboo. They are occasionally considered to be pests for this behavior, though preventing them from nesting in a deck or siding may be as simple as good upkeep (i.e. fresh paint, deck stains, et cetera). Tunnels extend around eight inches, maybe more, with around six or seven partitioned cells. Adult mothers may live two or three years, an oddity for a native bee. Occasionally several females, including a mother and a few of her daughters, will share a nest entrance and provision cells together thus the primitive eusociality. Woodpeckers and other predators and parasites feed on the contents of the cells.

Carpenter bee male, Xylocopa tabaniformis

Male carpenter bees are usually slightly smaller than females. The local species have males with light colored hairs on the thorax while females tend to be entirely black, including their eyes. Males have green eyes, and in my opinion are quite stunning. Males are also territorial and usually claim a patch of flowers as their own. They will chase and grapple with anything that enters their domain, including competing males, other types of bees, flies, falling leaves, pebbles thrown in their general direction, you get the idea. On occasion males even try to ward off humans who get too close, but one should not bother being intimidated because the males, like all male Hymenoptera, lack stingers.

Small carpenter bee, Ceratina sp.

Close relatives to the large carpenter bees, small carpenter bees (Ceratina spp.) are quite dissimilar to their large cousins. Many species of Ceratina are so small they are likely to be missed altogether, or mistaken for small flies. Rather than nesting in wood, they nest in broken pithy stems of woody plants, usually those in the Rosaceae (i.e. roses, blackberry). The small bees also have an affinity for flowers of plants in the rose family, though they visit a wide range of plants in many families.

Small carpenter bee, Ceratina sp., on Monardella villosa

Small carpenter bees are lightly haired, with sparse white scopae on the hind tibia. Many species are actually a dark metallic green, though by casual observation it's not always easy to notice. Females have bluntly pointed abdomens. Some species have white markings on the lower part of the face. Similar insects are other small bees, particularly masked bees (Hylaeus spp., Colletidae), and small aphid wasps (Pemphredoninae, Crabronidae).

Long-horned bee male, Melissodes sp.

Long-horned bees are named for the very long antennae of the males, as seen in the photos. The common name actually represents several genera in the Eucerini tribe including Eucera, Melissodes, Tetraloniella, and perhaps a few other less common genera as the males are all endowed with very long antennae. Melissodes, and others, nest in the ground. Species in my region in Southern Oregon are active in late summer. They are very fast and difficult for me to photograph, though this only adds to their allure. I caught and chilled two males for the purpose of identifying them, allowing them to warm up and fly away after I had taken a few photos.

Long-horned bee male, Melissodes sp.

Male Melissodes have two-toned antennae which are dark on top and buff colored on the bottom. They don't seem to be territorial, as many males and females are often found in a particular flower patch. In my observations males attempt to mate with females while they are foraging, and both males and females will attempt to mate repeatedly, if the males don't get bucked off by the females. I have not seen long-horned bees attracted to gardens, rather they seem to prefer specific patches of plants. I have observed them visiting Madia elegans in the middle of summer, normally in the morning before the heat sets in. The other plant I have seen them visit en masse is pennyroyal, Mentha pulegium, a nonnative from the Old World. I wonder if their floral preferences are influenced by the closeness of the floral resources to their nesting sites, often congregations of many nests in a relatively small area.

Digger bee, Anthophora urbana (2017)

A common digger bee, Anthophora urbana is native to much of the West. These fast flying bees are active for much of the growing season so long as there are floral resources to their liking available. I captured and chilled this female visiting vinegarweed, Trichostema lanceolatum (Lamiaceae). Like the long-horned bees, they collect dry pollen with scopae and nest in the ground. The genus Habropoda, related to Anthophora, are also found in this region but I have only observed them earlier in the year when the fruit trees are in bloom. These bees are able to sonicate, like the other large bees of the Apidae.

Cuckoo bee, Nomada sp.

The Apidae contains a vast group of upstanding citizens, those which contribute to pollination and demonstrate what we would call ethical guidelines. But what do bees know of ethics? A handful of bees in the Apidae are kleptoparasites, meaning they steal the nests of other bees for their own brood. These are known as the cuckoo bees. Kleptoparasitism is known to several bee families, and even within genera that are mostly not kleptoparasitic. The genus Ceratina, for instance, has a few kleptoparasitic species. The most notable feature of cuckoo bees is that females lack any mechanism for collecting or carrying pollen, as the nest they steal is already provisioned with the necessary pollen by the host bee. Cuckoo bees, thus, do not make a large contribution to the pollination effort. They are, anyway, interesting and beautiful, often wasp-like due to their lack of hair.

Cuckoo bee, Nomada sp.

Cuckoo bees enter the nests of other bees usually solitary bees like digger bees, and lay an egg in one or more cells prepared by the host. The cuckoo eggs typically hatch before the host egg, then the larva kills the host egg or host larva with a pair of over sized mandibles. Development the happens as expected, but what emerges from the host cell is not the host species, but the parasite. Fret not, for the presence of such bees indicate a strong population of host species, which is good overall.

Virgin honey bee queen

Are there more? Absolutely! There are over 1,000 species in fifty genera in North America, and well over 5,000 species worldwide in at least 200 genera. That is a lot of bees. So, when someone mentions bees it might be prudent to ask them to be more specific. The word bee is used colloquially to talk about honey bees, which do not accurately represent the family as a whole but are rather more of an oddity. Not to say that honey bees shouldn't be appreciated, but rather I think one should appreciate them for how they serve us and what they represent evolutionarily as the most advanced of all bees. I don't mean to say they are better than other bees, just different. I love all bees, and I think you should too. That is my message. (And wasps, now read this.)

(Photos taken with my trusty Nikon COOLPIX B700 and Samsung Galaxy S7 Edge. Some photos taken with the S7 through the eyepiece of a dissection microscope courtesy Old Sol Enterprises)

Daucus carota

noreply@blogger.com (Travis Owen) — Fri, 15 Sep 2017 16:00:00 +0000

Daucus carota with pollinators (2015)

Drive through rural Southern Oregon in summer, and you will see wild carrot blooming in dry pastures and on roadsides everywhere in low elevations. Wild carrot, progenitor of the cultivated carrot (D. carota ssp. sativus), is a naturalized nonnative originally introduced from Eurasia, the exact conditions of the introduction aren't clear. Carrot is biennial, meaning that the plants usually bloom the second year from seed, then produce their own seeds and die.

Daucus carota blooming in a dry field (2015)

One upon a time, long before European settlers first made this land their home, the flora was probably very different. The introduction of grazing livestock such as sheep or cattle created conditions that favored grasses, including many introduced grasses, after much of the native flora was grazed away. Compaction of the topsoil and soil disturbance, as well as the intentional and accidental introduction of a variety of nonnative plant species with a variety of uses were other factors that changed the landscape. These introduced species, as well as the physical impact settlers had on the land, changed the landscape in ways that are hard to imagine.

Each tiny floret will produce seed. In the very center of the umbel is one or two reddish to blackish colored florets that are thought to have some role in attracting pollinators, or at least that's the prevailing theory since there isn't any other obvious reason for it.

Small beetle and masked bee, Hylaeus sp. (2017)

But all is not lost. Wild carrot is a useful insectary plant with flowers that support a wide variety of pollinators and other beneficial insects. An umbel, the inflorescence in which the flowers are born, contains hundreds of tiny florets complete with pollen and easily accessible nectar. Following I will exhibit a few of the floral visitors I have observed on carrot umbels over the past few years.

Honeybees

Among the beekeeping community at large, there is an urban myth that honeybees only visit carrot flowers every ten years and that it causes the honey crop to become bitter and distasteful. While I can't attest to the latter, I can say without a doubt that I have observed honeybees working carrot flowers for three years running. This may be more indicative of the lack of available forage for honeybees than their actual interest in carrot as a resource. In commercial pollination of carrots for seed by honeybees, how much seed is set is dependent on the specific phenotypes as well as competition from other host plants in and surrounding the carrot crop. Native pollinators may be found to be more efficient and willing to visit carrot flowers, as illustrated by the series of photos below.

Masked bee, Hylaeus sp. (2017)

Small native bees, such as masked bees and small carpenter bees (Ceratina sp.) seem to like carrot flowers more than larger bees. While I have seen the occasional bumblebee visiting wild carrot flowers, they seem to be the exception to the rule. Small bees likely forage on wild carrot frequently where their habitat abounds, though their small size makes observation somewhat of a challenge. Perhaps a fine bug net swept through a dozen or so carrot umbels in bloom would yield a handful of small bees and other small pollinators that may otherwise remain unnoticed or unseen.

Crabronid wasp, Ectemnius sp. (2017)

In my observations, there has been a greater diversity of wasps visiting the flowers of carrots than bees, though I suspect this is purely a condition of the time and place of my observations. Crabronid wasps such as in the above photo were also less common than sphecid wasps in my observations. Crabronids are theorized to be the closest living relatives to modern bees.

Digger wasp, Sphex lucae (2017)

Digger wasps, such as Sphex, are often relatively large and in my opinion beautiful solitary wasps. Most wasps, in general, have mouth parts poorly equipped for foraging for nectar like bees, but have mouth parts well suited to accessing the nectar of plants in the Apiaceae, or parsley family, to which carrot belongs. This explains why many carrot relatives are excellent insectary plants. Sphex prey on katydids, which are paralyzed and buried in a subterranean nest for the larvae to feed on.

Thread-waist wasp, Ammophilinae (2015)

Thread-waist wasps, such as Ammophila, are common visitors to a variety of flowers. I postulate they have longer than average proboscises than some of their relatives since they visit flowers such as Allium and Echinacea which have nectar held deeper in the florets than Daucus.

Spider wasp, Pompilidae (2015)

Predatory spider wasps are some of my favorite types of wasps. They are solitary and often have underground nests like many sphecids and crabronids. As the name implies, spider wasps hunt spiders. A single spider, paralyzed by one to several stings, in dragged to the wasps nest where a single egg is adhered to the spiders underside. The wasp larvae feeds on the living spider.

Tiphiid wasp, Paratiphia sp. (2017)

Tiphiid wasps are normally parasitoids of soil dwelling beetle larvae, sometimes other wasp or bee larvae inhabiting soil or rotten wood. Tiphiids are hairy, the first time I saw one I thought it was a bee.

Tiphiid wasp posterior, kind of distinctive (2017)

Aphid wasps, Stigmus sp. (2017)

Solitary aphid wasps, Stigmus sp., were about the same size as masked bees and likewise almost completely devoid of hair of their bodies. When I took this photo I had mistaken it for Hylaeus, a masked bee. They are very small, and many features are lost to unaided eyes. That is one of the things that I love about macro photography, synonymous to a mobile action microscope. These small wasps nest in hollow twigs, and mass provision cells with aphids.

Potter wasp, Eumenes sp. (2017)

Potter and mason wasps (Eumeninae) are solitary members of the Vespidae, the family containing some of the more infamous members of wasphood (i.e. yellowjackets). Unlike their eusocial cousins, mason and potter wasps are not aggressive and very unlikely to sting. Those who have been stung by them report a disappointing experience, not for any pain the sting inflicted, but for the lack of anything notable. Yellowjackets and paper wasps evolved venoms that inflict not only more pain but more damage to tissues. This is because eusocial species must not only defend themselves but their nest mates, their nest, and all the young and resources within the nest. Solitary species invest relatively little into the construction of their nests, thus will readily abandon them and start anew rather than risking their lives defending the nest.

Vespid wasps, Polistes aurifer and Vespula pensylvanica (2015)

Eusocial vespids also visit Daucus. The only local eusocial vespid I have not seen visiting carrot flowers are the large bald-faced hornets, Dolichovespula maculata.

Polistes dominula (2015)

The majority of wasps don't collect pollen for their young, and most don't collect nectar either. Adults of many species will visit flowers for nectar for their own use, fight takes a lot of energy and nectar is a good source of carbohydrates. Some wasps do collect and store a small amount of nectar, a sort of wasp honey. I have seen this substance in Polistes wasp nests of a few species, and have heard that yellowjackets do this too though their nests are harder to observe since they are usually underground.

Syrphid fly, Myathropa florea (2017)

A variety of wasp or bee mimics are attracted to the small plentiful flowers of carrot. Various hoverflies are commonplace, many of them being wasp/bee mimics. They fill several roles as pollinators, predators (larvae), and recyclers. The species photographed above feeds on bacteria in the larval stage within rotting logs or other rotting plant matter. Their color patterns are an aposematic defence, a form of Batesian mimicry. Flies don't have stingers, and all but a few (no syrphids) can bite.

Tachinid fly, Archytas sp. (2015)

Tachinid flies, like syrphids, are good to have around. Most are parasitoids, the larvae feeding on an array of pest species such as moth larvae or other flies.

Ornate checkered beetle, Trichodes ornatus (2015)

Mimicry isn't restricted to flies. Beetles also employ color schemes which might give pause to potential predators. The ornate checkered beetle is often found on flowers. Adults take nectar, and may feed on pollen as well. They are parasitoids of ground nesting bees and some wasps.

A mating pair of blister beetles, Epicauta sp. (2015)

Blister beetles are numerous in late summer, often seen on flowers in dry areas. They produce a chemical compound that, when touched by human hands, cause blisters to form. Adults eat flower parts and perhaps take some nectar. They are common where Madia elegans is in bloom, in my experience.

7-spot ladybird beetle (2015)

Ladybird beetles, like the honeybee, are one of the poster children representing beneficial insects and often adored by young children. Like the honeybee, the seven-spot ladybird beetle is an introduced species sold for aphid control. There is anecdotal evidence suggesting that they negatively impact the native species.

By the first frost in fall, most of the fruits have fallen onto the ground awaiting amenable conditions in which to sprout. Many of the plants in the Apiaceae have seeds which require light to germinate, so when sowing it is best to just cast them out onto a prepared seed bed and tamp them in. Many also benefit from a cold treatment, so a fall sowing is beneficial. These tips I have learned from observing how the plants sow themselves. By late fall, most of the seeds of dill, carrot, parsley, lovage, angelica, and nearly all others have naturally been dispersed by some means. Most also have taproots, so direct sowing is the easiest way to grow them.

When the seeds don't fall, they are happy to germinate right on the inflorescence. This is a futile endeavor, of course, and as soon as the rains cease in late spring they are doomed.

Thoughts on Rabbitbrush and Other Things

noreply@blogger.com (Travis Owen) — Wed, 13 Sep 2017 01:52:00 +0000

I’ve recently been jonesing to get out and observe rabbitbrush out in the wild and photograph the pollinators that come to visit its flowers. It’s an interesting plant, and one of the latest natives to bloom in dry rocky sites in the Rogue Valley. Rabbitbrush (Ericameria nauseosa) is a grey-greenish shrub in the sunflower family (Asteraceae) with smallish yellow florets born in a panicle and bearing no resemblance to sunflowers or daisies. The plants are prodigious nectar producers, and attract a wide range of pollinators throughout their range, often attracting any and all of whatever pollinators are still active in the area at the time. So I was on a hunt to find these plants and get my fix. Being short on time, I went somewhere not too far from where I live, out past Emigrant Lake in Ashland. I saw a few of the shrubs on the side of the road, but curiosity and the urge to explore took the wheel and I kept driving way out past the lake to wherever Oregon Route 66 would take me.

Eristalis sp.

I pulled over at a small turnout where I saw a fair amount of rabbitbrush in bloom and proceeded to observe. The plants like to grow in places that seem inhospitable to many other plants, such as on the sides of rocky bluffs or, in this case, on the steep slope on the downward side of the road. I slid several times, my shoes and socks filling with gravel and dirt, but fortunately stopped myself from cascading down the bank, the bottom of which was far out of reach.

Paradejeania sp.

Last year I had photographed a variety of wasps and honeybees on rabbitbrush blooming along Emigrant Lake, but out in the hills, miles and miles out on OR-66, mostly syrphids (Eristalis spp.) and tachinid flies (Paradejeania sp. and others) were to be seen visiting the flowers. A single thread-wasted wasp and a couple small sweat bees were the exceptions. Bald faced hornets were hovering around the bushes, probably seeking an unfortunate insect of sorts to bring back to the nest, and having no interest in the flowers themselves.

Rabbitbrush with a fly gall, Aciurina bigeloviae

I moved on, my yearning to explore now taking full control of the situation, this time diverging onto a gravel road with a sign that read Hobart Buff Trailhead. I was entering the Cascade-Siskiyou National Monument. Why not? I know nothing of this particular place, yet I have wanderlust like a bird has feathers so it wasn’t even a question as to whether or not I should try it out. But I was also looking for flowers, a futile goal since by late August most of the native wildflowers are spent and most of the native pollinators won’t be seen again until next year. Why should I grasp for something that I just know isn’t there, I wondered? Surely there are other reasons to go on a hike, such as health, spontaneous experience, fun, et cetera.

Centaurea sp. with a fly

I stopped midway somewhere on the gravel road between the 66 and the trailhead to investigate some flowers in bloom. Weeds, nonnatives, were the predominant plants in this clearing. A white flowered Centaurea was the most prolific, and although the sun was shifting ever closer down towards the horizon there were still bumblebees on the flowers, as well as a small assortment of flies.

Still smokey on the PCT

Upon arriving at the trailhead, I was greeted by a peculiar hissing sound coming from behind my truck. I was thrilled to discover the rough road had torn a small hole in my tire. After gleefully replacing the leaking tire for my spare (I was not excited about the thought of waiting until after I'd hiked to change it), I was off on my merry way down the trail, camera and walking stick in hand.

Despite my initial disappointment beauty and delectation were there to be found. There was practically nothing in bloom on the trail, except for one tiny tiny species with minuscule flowers, hugging the ground and ready to be completely unnoticed by any reasonable hikers. What were once wildflowers a month or two ago were now crispy dry remnants with ripening seeds or already dispersed seeds apparent by open and empty seed capsules.

The maturation of seeds and seed dispersal facilitated by uniquely evolved seed carrying structures is of great importance. Incidentally, seeds and seed capsules are quite interesting. Plant life, as well as plant death at the end of an annual cycle, is beautiful.

I was pleased with what I had seen, though I had to shift my goal from finding flowers and pollinators to finding other beauty. I enjoyed challenging myself to find other things to appreciate, a task often easier said than done in my experience. My mind was unsettled, and thoughts drifted to a region of my mind neither pleasant nor unpleasant.

I have been reading a lot of the works of Albert Camus this year, philosopher and author from the first half of the 20th century. He was considered an existentialist, with some similar themes as found in the works of Nietzsche, Kierkegaard, and Sartre. Most recently, and perhaps appropriately in the midst of the thick isolating smoke we have experienced this summer up to the last week or so, I have been reading The Plague. In this book Camus depicts a plague stricken township completely from the outside world via a strict quarantine. The epidemic lasts for a very long time, around a year if I understood correctly, and is dubbed "a long period of exile" by the narrator. Whether one tried to escape the city, an analogy to trying to escape death, or attempts to aid in some way such as taking care of the ill or any other noble cause, all are on the same figurative boat and will all face the same fate, be it from the bubonic plague or any other cause. In Camus' view, even though taking the noble route resulted in the same as any other route, it was still worthwhile and thus meaningful. This is what I thought about during the hike.

I agree with Camus in that everything is only temporary, be it anything deemed good or bad in life, exemplified by flowers and the short lives of bees and other pollinators. I don't think bees care about flowers or their survival. This doesn't mean they don't need flowers. In this way we aren't that different. We have needs of various sorts but often don't care about them unless we choose to. I believe everything comes down to choice. I am also a relativist and believe much of our choices are biased, subconsciously tainted from our families, experiences, cultures, et cetera. How much is nature and how much is nurture is uncertain, and I don't know if there would be a way to test this without bias.

One thing that draws me to flowers and pollinators is that they just do without wondering if it is right. Their actions (not choices, I am hesitant to believe flowers or pollinators have free will) are purely for survival, and fitness, and not about contemplating their lives. They have an advantage, not burdened by choice. But with choice comes great power, great opportunity. But is this a good thing? I am sometimes inclined to think not, for reasons that are far too many. But hey, look at these photos of blister beetles eating the flowers of Madia elegans!

Madia elegans is one of y favorite native plants. They are peculiar. The flowers open at dusk close in the morning. The entire plant is covered in sticky, resinous hairs that simultaneously reduce desiccation from the sun and trap small insects that may cause the plants harm. And they grow like weeds, in the best possible way.

Life has me trapped in a continuous cycle of sleeping, eating, and earning money to redeem for basic survival needs. Life is not always pleasant, and it is not always a choice to think otherwise. I choose to embrace nature in any way that is within my power, because it brings me deep satisfaction. I don't know if I'll ever make a difference beyond my own life, but really this shouldn't matter. We should choose to be happy for our own sake, because doing otherwise is absurd. Enjoy each day, even if it's for small simple pleasures, like a sunset. I live a life free from many of the horrors many people on this earth experience on a daily basis. If I don't appreciate that, then I am truly living in absurdity and life really is meaningless.

Trichostema lanceolatum

noreply@blogger.com (Travis Owen) — Fri, 01 Sep 2017 14:07:00 +0000

Trichostema lanceolatum is visited by woodland skippers, Ochlodes sylvanoides. (2016)

Most plants, without question, are first seen, but there are also those that are first smelled. While taking a walk through a local woodland clearing one day, a sudden strong scent of gasoline or turpentine wafted into my face. It had perplexed me for some time, until I discovered that the origin of the scent was from a plant. I have come to look for this plant every year in late summer and fall, as it is one of the last native plants to bloom in dry areas in the Rogue Valley of Southern Oregon.

Masked bee, Hylaeus sp., coming in for a landing (2017)

Trichostema lanceolatum Benth. (Lamiaceae), or turpentine weed, is an annual Western native from California north into Washington. The plants grow to a height of up to ten inches, often branching from the base. The leaves are of a narrow and tapering to a point at the tip (hence lanceolatum, or lance shaped). The entire plant is covered in dense hairs (trichomes) that secrete a resinous oil, the source of the strong scent which I think is more akin to gasoline than turpentine (or camphor, as the common name camphor weed suggests).

Trichostema lanceolatum (2016)

The fascinating flowers of turpentine weed are one of the features that make this genus so interesting to me. A related species, Trichostema lanatum, are known as blue curls, presumably named for the flowers. The anthers and stigma are arched out away from the flower, awaiting contact with a pollinator. The shape of the flower, with the arching reproductive structures, are similar to others in the Lamiaceae such as rosemary, Rosmarinus officinalis.

Trichostema lanceolatum with seed pods (2015)

Flowers are borne on a type of inflorescence called a cyme. Multiple flowers bloom at once, maturing from bottom to top. Each flower produces up to four nutlets which fall and germinate the following Spring. They seem to respond well to mowing, or perhaps mowing allowed me to spot them easier when I first discovered them in bloom. I also speculate they may have benefited from wildfires in the ages before European settlers inhibited the annual fire cycle.

Trichostema lanceolatum (2015)

Visible here is the flower structure, the anthers have yet to fully expand, or they have been damaged in some way and are deformed. You can also see the dense hairs, possibly serving multiple purposes. As seen in Madia elegans, the resinous hairs (or trichomes) probably help the plants retain moisture during the heat of Summer. Another possibility is that the hairs, or perhaps more importantly the potent resin, somehow protects the plant from certain insect pests.

Trichostema lanceolatum variability (2015)

Those who are color blind may have some difficulty finding the flowers in this image, and if the reader is color blind I would appreciate some input on this. Mule deer, important herbivores in the Rogue Valley, are red-green color blind like some humans. I have long hypothesized that the color of some plants that bloom in dry grass-dominated areas in summer help prevent them from herbivores when the grass is allowed to remain. This is, however, purely speculation. The scent of these plants may be so strong that the smell alone may deter herbivores. I have never seen damage from these plants from herbivores, at any rate.

Trichostema lanceolatum (2015)

Many of the plants lean generally towards the South, the flowers all being arranged more or less on the same side. For perspective, the South is where the sun is as the days shorten. In winter, the southern exposed sites are the only places to receive a full day of sun, while exclusively northern exposed sites are permanently shaded through winter's entirety. Seed survival through winter, I speculate, may also be dependent on a southern exposure, while a northern exposure has historically resulted in soil that is cold and frozen solid for winters entirety and in general is not very kind to flower seeds (personal observation).

Trichostema lanceolatum (2015)

The habitat is best described as hot, with what is suspected to be nutrient deficient soil, possibly serpentine (though I do not believe this plant is an indicator of serpentine). The biochemicals secreted by this species and others have allelopathic properties, reducing competition. This could explain why the areas I have found them growing are so sparsely vegetated while the surrounding area is comparatively lush. Growing in poor soil is also a technique some plants utilize since there is generally less competition from other plants.

Trichostema lanceolatum (2015)

I have observed few native blooming plants in the area at this time of year, limited to Madia elegans, Achillea millefolium, and Eschscholzia californica, aside from the Trichostema. The most common flowering plant here, seen in fields everywhere, is the invasive European native Daucus carota, the wild carrot. This leads me to ask the question, is that all that native pollinators had to forage on? Were there once other wildflowers here for native bees, or is the near end of the pollinator season naturally deficient in native wildflowers, corresponding to lowered numbers of active native pollinators late in the growing season?

Pollinators on T. lanceolatum (2016)

Perhaps before European settlers made this their home, the few native plants that were blooming were more widespread. The introduction of nonnative grasses and the overgrazing of livestock has probably dramatically and devastatingly altered the landscape to the detriment of biodiversity. What may now look like a native grassland with four or five native species of wildflowers may have once harbored dozens more. We can not know what has been lost, and I am reluctant to believe it can ever be restored.

Ochlodes sylvanoides (2016)

The most common pollinators to visit vinegar weed have without question been woodland skipper butterflies, Ochlodes sylvanoides. Woodland skippers were numerous and frequently visited many flowers on each plant and on multiple plants, suggesting they may be the primary pollinators of T. lanceolatum in this region. I have spent hours observing these plants in different locations over the past three years, and the woodland skippers were and still are the most frequent and numerous pollinators to visit.

Ochlodes sylvanoides (2016)

An interesting characteristic of the flowers of Trichostema lanceolatum is that the reproductive organs swing down onto the body of the pollinator when they land on the flower, causing pollen to be transferred. When the pollinator departs, the stamens and stigma return to their original position.

Ochlodes sylvanoides (2016)

The woodland skipper is active in the fall. Being grass skippers (Hesperiinae) they use host plants in the grass family (Poaceae) including Bermuda grass (Cynodon dactylon), canary grass (Phalaris sp.), wildrye (Elymus sp.), and wheatgrass (Agropyron sp.) though that is not a complete list. They are native to the Western US as far east as North Dakota and north from British Columbia to Saskatchewan. During their relatively brief adult lives, O. sylvanoides mates, imbibes nectar, and lays eggs which hatch in a matter of weeks. First instar caterpillars hibernate through winter, complete their feeding in the spring, diapause in summer as fully grown caterpillars, then build their chrysalis and pupate til emergence in the fall.¹

Ochlodes sylvanoides with pollen on the thorax (2016)

The woodland skippers thorax, in my observations, made contact with the reproductive parts of the majority of the Trichostema lanceolatum flowers they alighted on. In a few instances when the skippers were at rest I was able to observe more closely and noted pollen on the throax of most, as seen in this photo. Crossing of pollen from one plant to the next is a highly likely scenario considering the rate at which these butterflies moved between plants.

Masked bee, Hylaeus sp. ♀ (2017)

Multitudes of small bees visit vinegar weed, though I have not observed honeybees to visit them. The most numerous bees have been small wasp-like masked bees (Hyaeus sp., Colletidae). They alight on the anthers, and also on the labellum, seeking either pollen or nectar, respectively. Masked bees are not considered to be very efficient pollinators, despite being bees. This is due to the scarcity of hairs on their bodies. Pollen is eaten, held in a crop, and regurgitated in a nest cell. Many other bees transport dry pollen on special hairs on various parts of their bodies called scopae. Some apid bees (honeybees and bumblebees), in contrast, transport pollen moistened by nectar in specialized structures called corbicula, patches of specialized curved hairs on the tibia of the hind legs of females.

Masked bees often nest in pithy stems or twigs, or occasionally in cavities in wood, or occasionally in soil, depending on the species. The walls of the cells are lined with a silk-like polymer which serves some role in protecting the delicate larvae from parasitic or microbial pests (Espelie et al. 1992).

Sweat bee, Halictus sp. ♀ (2017)

Slightly larger than the masked bees, sweat bees were also observed visiting vinegar weed. Unlike the masked bees who would land on different parts of the flowers, the sweat bees nearly always landed on the anthers in my observations. Halictid bees are typically ground nesting bees, nesting in excavated burrows in bare ground. Some species, as well as the related Lasioglossum, are primitively eusocial or communal. In extreme cases this means that there may be one or a few dominant laying females while the rest are essentially subordinate workers though they may also be fertile. Alternatively they may share only a nest entrance while each female excavates and provisions her own cells within. Many still are completely solitary, though of those some may nest in congregations.

Digger bee, Anthophora urbana ♀ (2017)

The largest bees I have observed visiting vinegar weed have been digger bees. I caught one very fast flying individual in order to identify it. Anthephora urbana are slightly smaller than honeybee workers. They are solitary ground nesting bees, with long tibial spines used for maneuvering inside the burrow and stabilizing the bee while excavating the cells. Digger bees are in the Apidae, the family of honeybees and bumblebees, and like bumblebees can sonicate while visiting flowers. Sonication, or buzz pollination, is when the bee vibrates its wing muscles while on a flower which causes the pollen to burst explosively from the anthers and due to a phenomenon which causes the pollen to stick to the bees body with electrostatic attraction. Flowers, thus pollen, are negatively charged due to being, quite literally, well grounded. Bees, and all flying insects, develop a positive charge.

Trichostema lanceolatum (2016)

I have received the testimonies from others that bumblebees or perhaps large carpenter bees also visit the flowers of vinegar weed, and appear to fit morphologically. I don't doubt their effectiveness as pollinators, in fact the "buzz pollination" of large native bees is highly effective for the pollination of many natives (particularly those with very sticky pollen, released only by the intense vibrating of the wing muscles). But I did not witness any large bees visit the flowers during my initial observations. Such is the variability of time and place. I have also observed beewolves, Philanthus sp., in the vicinity of vinegar weed though I have not observed them visiting flowers. Beewolves, predatory wasps in the family Crabronidae, have common ancestors to modern bees.

Trichostema lanceolatum (2015)

Plants like this, plants which may be easily overlooked or may even evoke amusing faces when they are detected by scent but unseen, are a particular area of interest for me. In fact, I am often enamored by the small and overlooked, including bees and other pollinators. These small creatures represent small personal discoveries which ignite my wonder and curiosity for the natural world, and inform my opinions of life as a whole. Small creatures with no voice of their own deserve to thrive just as much as I do, yet I dread that their lives are hanging by a thread. Many people don't value life beyond their own. Whether this is resultant of the belief that people are superior to all other lifeforms or the thought has simply never occurred to them is unclear. But small life inspires me, and my goal is that by writing this and taking photos this life may also inspire others.

Trichostema lanceolatum (2015)

Mount McLoughlin

noreply@blogger.com (Travis Owen) — Sat, 12 Aug 2017 16:43:00 +0000

I recently took my first hike up Mount McLoughlin, the tenth highest mountain in Oregon at 2895m (9,499ft), located in the Sky Lakes Wilderness between Medford and Klamath Falls. It was hot out, the forecast said it was going to be sunny, around 100°, and dry. I drove there on highway 140 with the windows down, letting the wind hit my face. I enjoyed the smells of dry meadows and pine trees as I took a few last glances at Mount McLoughlin before it went out of sight.

The map at the Summit Trail trailhead

From Medford, it takes a little over an hour to reach the summit trailhead, but shortly after setting out I lost sight of the mountain as I entered the woods and foothills surrounding the mountain. Moving ever closer to the mountain the only flowers I could see were fields of wild carrot, Daucus carota, an introduced species from Europe and southern Asia. Occasionally I spotted common chicory, Cichorium intybus, another introduced species hailing from Europe, in the occasional roadside ditch.

Fireweed, Chamaenerion angustifolium

Driving onward, and subsequently higher in elevation as I wound through the Rogue River-Siskiyou National Forest on the 140, more wildflowers came into view. Fireweed was one of the most conspicuous, as its name not only describes an aspect of its biology, but also evokes its vibrancy and large stature. As the name suggests, it often appears after wildfire where it is native, when competition is burnt to the ground.

Henderson's checkermallow, Sidalcea hendersonii

Another vibrant Oregon native, named after plant explorer and collector Louis F. Henderson (1853-1942), Henderson's checkermallow is related to hollyhocks and grows much like a miniature hollyhock, growing often to around two feet in height. Henderson is the namesake of some of my favorite wildflowers, including Dodecatheon hendersonii, Erythronium hendersonii, and Triteleia hendersonii, all native to this region.

Californioa goldenrod, Solidago velutina ssp. californica

Gracing the parking area at the Summit Trail trailhead, California goldenrod grows to around sixteen inches. Topping each plant is a pyramidal panicle of tiny yellow daisies. They seem to like sunny edges of forest clearings, or in this case, roadsides bordered by a coniferous woodland.

Twinflower, Linnaea borealis, and a bumblebee

I had arrived at the trailhead and was ready to embark. Near the parking area is a cool shady area with abundant shade loving wildflowers such as Western Prince's pine (Chimaphila umbellata) and twinflower. The small honeysuckle relative, twinflower, so named because they nearly always have flowers in pairs. Small bumblebee workers, unfortunately unidentified, were vociferously visiting each flower yet in a manner that made them difficult to photograph by this amateur, thus the blur.

Western Prince's pine, Chimaphila umbellata

I was quite stunned by the beautiful yet diminutive Chimaphila umbellata, a relative to blueberries and azaleas. This plant is categorized as a shrub, though the individual plants measured less than six inches and had an even smaller footprint. Small pendant flowers, maybe half an inch wide, look towards the ground awaiting a pollinator. Curiously I saw not a single pollinator visit these flowers, even though they were the most abundant wildflower on much of the lower forested portions of the hike. Intrigued, I had to smell one, and it had a musky scent similar to that of Prunus or Ceanothus blossoms but reduced in intensity. Peculiar.

Rough hedgenettle, Stachys rigida

Approaching the Cascade Canal that runs parallel to the road to the trailhead, I noticed the nuanced flight of bees, and on closer inspection (by which I mean navigating some rocks to the center and avoiding falling in the water, refreshing as that may be) it was revealed that bees they were! A pair of bumblebees, regretfully unidentified, were foraging on the mint in a small island in the canal. The mint is rough hedgenettle, a wild relative to lambs ear (Stachys byzantina).

Scarlet gilia, Ipomopsis aggregata

After crossing the bridge that spans the Cascade Canal, there was a sunny clearing with a variety of wildflowers, and hordes of butterflies.

Into the woods

The next phase of the trail was through the woods. The ground was mostly flat, and the air was refreshingly cool. Giants loomed above me, mostly red fir (Abies magnifica), and wildflowers dotted the landscape along with woody debris. I exchanged pleasant greetings with passing hikers, most of which were returning from the summit. I had not thought to find out how long this hike would take, until by chance I asked some hikers who informed me that the summit was many hours away. There was a moment of discouragement which I hastily disregarded and continued down the path towards the summit.

Scouler's harebell, Campanula scouleri

The tiny Scouler's harebell was in bloom near the trail along with other diminutive wildflowers. The similar but taller Campanula prenanthoides is visited primarily by bumblebees (personal observation) and I would expect that C. scouleri to be the same, though there was a notable lack of visible pollinators at the forest floor at the time of my being there.

One-sided wintergreen, Orthilia secunda

One plant that intrigued me was the aptly named one-sided wintergreen. Aptly named, because the flowers all hung on a single side of the scape, though in retrospect I regret not crushing and sniffing the leaves to see if they smelled minty, but who does that really? It was the second most common plant in bloom after the Chimaphila, at least in the forest at the low end of the trail. The flowers are green, urn shaped, with long stigmas extending out. I found them to be charming.

Hadromyia pulchra male

Once under the cover of the trees, there was only the occasional buzz of a flying insect (not counting the mosquitoes which did not bother me too much, but, as I was informed, "mauled" some of the other hikers). This large syrphid fly buzzed very loudly indeed, so much so that I initially mistook it for a yellowjacket. I think this was a case of Batesian mimicry in which the fly, harmless, makes itself as conspicuous as possible by flying right up to and around a potential predator, mimicking the sound of a wasp, and even resembling a bee or wasp in the color scheme.

Laphria ventralis male

A large male robber fly perched on a rotting log, seeks a mate, a meal, or both.

Carpenter ant, Camponotus sp.

The most conspicuous insects of all were the carpenter ants. They are large wood nesting ants that live in dead, often rotten, wood. They are omnivorous, eating both dead and living insects, honeydew, and occasionally nectar. They don't eat wood, but rather remove it from their nest as they create tunnels and chambers for food and larvae.

Townsend's chipmunk, Neotamias townsendii

Chipmunks and birds were the extent of the vertebrate wildlife I saw. Birds were mostly heard but not seen. Chipmunks have a strange call, and I've often mistaken the call of squirrels and chipmunks for bird calls. Caught this one red handed.

White-flowered hawkweed, Hieracium albiflorum

Common, and probably overlooked or underappreciated, white-flowered hawkweed flowers by the edge of the trail. Each small yellow stigma is part of a single flower, the entire daisy is in fact an inflorescence.

White-veined pyrola, Pyrola picta

White-veined pyrola, very similar superficially to some species of Goodyera, except the venation is quite different. Goodyera is an orchid (Orchidaceae), but Pyrola picta is in the Ericaceae, along with Chimaphila umbellata and Orthilia secunda.

Spotted coralroot, Corallorhiza maculata

A variety of parasitic plants grow along the trail, yet the only thing they seem to have in common is they lack the ability to photosynthesize and instead rely on other sources for nutrients. Corallorhiza is in the orchid family, and rely entirely on mycorrhizal fungi. It seems unclear whether coralroot is insect-pollinated or self-pollinated.

Spotted coralroot, Corallorhiza maculata

Small spots on the lip of the flower suggests that insects are the primary pollinators of coralroot, though in the absence of reliable pollinators the plant has evolved a backup strategy to pollinate itself and form seed independent of insects. Cross pollination is preferable for most plants as it diversifies the gene pool, though some plants are capable of selfing without suffering from the effects of inbreeding.

Striped coralroot, Corallorhiza striata, going to seed

Dutchman's pipe, Monotropa hypopitys

Unsurprisingly, another ericaceous plant, this time in the form of a parasitic plant that does not produce chlorophyll but rather survives through the mycorrhizal network.

Dutchman's pipe, Monotropa hypopitys, close-up of flowers

Gomphaceae (?)

Fungi are ever important in the forest, and everywhere else. Mycorrhizal fungi, those which form mutualisms on and within plant roots, are thought to have relationships with the majority of the plants on the Earth. Mutualistic, because both parties benefit from the relationship. The fungi feed on root exudates, and the plant benefits in multiple ways from the expanded surface area of the root system as the fungi attain a greater amount of water and nutrients than the roots can do alone, and the fungi block passage of pathogens, among other things.

Nearing the subalpine zone, many broken and stunted trees become apparent. The trail at this point is arduous at best, an uneven winding stairway of large angular volcanic boulders of unequal size.

The trail, at this point, became very difficult for me. But, turning around periodically and forcing myself to stop was motivating because I got to see my progress. Passing hikers informed me that the summit was still at least two or three hours away.

Western bleeding heart, Dicentra formosa

Entering the subalpine zone, the forest opened up and I began to see flowers again. Trees were not as tall, and not as frequent. The air, even in the direct sun, felt cool. A light breeze was invigorating. I wanted to keep going. My body said no while my mind said yes. I was inspired most of all by this single plant, the bleeding heart fed my curiosity and I wanted to see what else was flowering.

Western bleeding heart, Dicentra formosa

Though this is a common wildflower, blooming months earlier in the lower elevations, this is the first one I've seen growing wild. The flowers are odd, pollinated by bumblebees. Dicentra is in the poppy family, Papaveraceae.

Western columbine, Aquilegia formosa

The higher one goes, the most compressed the flowering season. Flowers that may bloom in spring in low elevations will bloom in summer in higher elevations. Western columbine, sharing the specific epithet with Western bleeding heart, is typically found blooming in the spring in similar niches as the bleeding heart in the lowlands. While both typically grow wild in somewhat shady, moist areas, in the higher elevations they grow in the open among rocks in full sun.

Western columbine, Aquilegia formosa

Columbines are bee-pollinated, though garden varieties seem to be subject to nectar robbing. Nectar is held within the spurs, and in gardens, in my observations, carpenter bees and bumblebees will just chew holes in the spurs to reach the nectar. Smaller bees were seen visiting the native columbines in both the lowlands earlier in the year and here in the subalpine environment.

Pinemat manzanita, Arctostaphylos nevadensis

Pinemat manzanita was frequently encountered spreading across stretches of the open area between the sparsely treed subalpine zone, sprawling over boulders. Most were forming berries, and only the shrubs higher up the mountain were in flower.

Cuckoo wasp, Chrysididae

Above the reign of the pinemat manzanita, boulders began to dominate the landscape between the occasional conifer. Between these boulders, pockets of soil has formed. In this soil, solitary bees and wasps were building and provisioning nests. One such wasp, a metallic green cuckoo wasp, was carefully visiting every nook and cranny for a nest entrance of another bee or wasp in order to deposit an egg in. Cuckoo wasps are stingless wasps whose larvae feed on the provisions of other wasps or bees, and thus they don't bother constructing or provisioning nests of their own.

Davidson's penstemon, Penstemon davidsonii

Rock plants, those cherished plants that grow in higher elevations among rocks, are one of the main reasons I wanted to make this hike happen. Davidson's penstemon is a low lying evergreen woody perennial that likes sharp drainage and rocky soils, clearly. Alpine and subalpine soils are formed very slowly, often composed largely of minerals and small quantities of organic mater that has deposited between rocks and in cracks by way of wind or from the sparse vegetation.

Davidson's penstemon, Penstemon davidsonii, with a small bee (Halictidae)

A small bee, a halictid bee of some sort, was foraging on the penstemon while I was admiring it. Small bees have the advantage of fitting inside flowers of this sort. The long floral tube of this penstemon may inhibit larger bees from accessing the nectar. The genus Penstemon, commonly called beardtongue, because of hairs within the corolla. The hairs, I think, serve to exclude inefficient pollinators or nectar thieves and perhaps also slows evaporation.

Davidson's penstemon, Penstemon davidsonii, with a small bee (Halictidae) inside the floral tube

California tortoiseshell, Nymphalis californica, on subalpine fir, Abies lasiocarpa

Both tortoiseshell butterflies and fir trees were very common on the mountain. The former were seen at all elevations, from the trailhead to the ridge. Their caterpillars feed of Ceanothus, their primary host, while the adults visit flowers. I also wonder if the adults feed on tree sap. Tree sap isn't an odd food for butterflies, some in the same genus feed nearly exclusively on sap, id est the Mourning cloak, Nymphalis antiopa.

Marum-leaved buckwheat, Eriogonum marifolium, male flowers

Upon entering the Upper Realm of the mountain, I saw more and more rock plants. Plants in this zone are mostly compact and small, an advantageous evolutionary trait. Conditions can be harsh at the top of a mountain, with conditions favoring desiccation. Plants must be able to survive long dry periods under snow, dry because all of the water is locked up in the form of snow until it starts to melt sometime in late spring or early summer. Growth tends to be very slow, the season is short.

Marum-leaved buckwheat, Eriogonum marifolium, male plant

Eriogonum, sometimes called wild buckwheat, was all over the place. Marum-leaved buckwheat, is interesting because the plants are either male or female. Male plants have yellow flowers, while female flowers are red and yellow (see below).

Marum-leaved buckwheat, Eriogonum marifolium, female flowers

In my experience eriogonums are visited by a huge range of pollinators, from many different types of bees and wasps, to beetles, butterflies, and various flies. Eriogonum marifolium is also a host plant for the buckwheat blue butterfly, Euphilotes battoides.

Marum-leaved buckwheat, Eriogonum marifolium, female plant

Marum-leaved buckwheat is native to the West, mostly Northern California, Southern Oregon, and maybe Nevada. Oddly, the common name comes from a similarity between its leaves and the leaves of Teucrium marum, a Mediterranean native, to which it isn't remotely related.

Oarleaf buckwheat, Eriogonum pyrolifolium

Sometimes plants are given common names that reflect their shapes or textures. Oarleaf buckwheat is so called because of the shape of the leaves. Another, less flattering name, is dirty socks, which is apparently in reference to its odor. Unfortunately, or fortunately, I can't confirm this.

Oarleaf buckwheat, Eriogonum pyrolifolium

Whitebark pine, Pinus albicaulis

Many of the trees were either stunted or contorted. Whether this is their normal growth habit, or a result of their growing conditions, or both, is unknown to me.

Thread-waisted wasp, Ammophila sp.

The compacted sandy soil of the trail is a popular nesting place for a variety of solitary wasps, and probably bees as well. Ammophila excavate nests in sandy soil. A female will first dig a nest then use a pebble and sand to conceal the entrance before embarking on the task of finding a caterpillar host. Once paralyzing a host and returning to the nest, she will reopen the nest and drag the caterpillar inside. Once enough hosts are delivered into the nest, or a single large caterpillar host is delivered, the female will lay an egg and seal the nest again with a pebble and cover it in sand to hide it. If the host caterpillar is large, the female wasp will be unable to fly with it and must drag the prey back to the nest, occasionally at incredible distances from the nest site. How they can find their way back to the nest whilst dragging their prey is a mystery. It would be similar to taking a plane from a house you just moved into and then walking backwards back home.

Kleptoparasitic pompilid wasp, Evagetes sp.

Spider wasps were everywhere if one was looking. Spider wasps, as the name implies, seek out spiders to provision their underground nests. A single spider is paralyzed, pulled into the nest (or sometimes in the spiders own burrow) and a single egg is laid upon it. The nest is then sealed, and the wasp larvae feeds on the living but immobilized spider.

Kleptoparasitic pompilid wasp, Evagetes sp.

Not all spider wasps capture their own spiders, such as the genus Evagetes. Instead, some steal nests from other spider wasps, kill the host larvae, and lay their own egg inside. This is called kleptoparasitism.

Square-headed wasp, subtribe Crabronina

Other wasps were certainly present, such as crabronid wasps. Solitary predatory wasps often visit flowers as adults, while capturing various prey insects to feed to larvae in nests in the ground, in wood, or in cracks and crevices between rocks. Bees are thought to have evolved from crabronid wasps.

Partridgefoot, Luetkea pectinata

I was running low on water, poor planning on my part, so I made the risky judgement call to leave the trail and cut across the scree to reach the snow patch on the southern side of the slope. I knew that if I could reach the ridge, I would find my way, and to reach the ridge all ?I had to do was climb up. I would not recommend anyone leave the trail on purpose. The trail becomes difficult to see anyway as it is mostly boulders, and footprints of previous hikers often leave the trail since they too sometimes take the wrong route.

Partridgefoot, Luetkea pectinata

The scree was very difficult to navigate, but I needed water. The sharp sand scattered with stones is held up by practically nothing, much like walking on a sand dune. Many plants thrive in this sandy medium. I needed to stop several times while crossing the scree, practically collapsing in a heap in front of the beautiful partridgefoot. There were a half dozen or so pollinators, flies and wasps I think, that quickly departed when I arrived.

The roots of subalpine and alpine plants are astoundingly long and deep. On the southern slope there were many plants pulled out of the ground, I imagine this was the work of heavy snow slowly sliding downhill. The roots need to reach deep in the sand to anchor the plants, and to find water. Water is not readily available in a scree, something that differentiates it from a moraine which has running water beneath it.

I had reached the snow, and took a long break before climbing up. I got brain freeze, an unexpected holdup. As I was sitting, I heard people calling out. I thought they were yelling to me, but then I heard a call from the opposite side of the slope. Someone had begun to descend the slope in the completely wrong direction. Fortunately his friends set him straight. I imagine a handful of people get lost up here every year.

Thick-leaved groundsel, Senecio crassulus

Castilleja sp.

Cobwebby paintbrush, Castilleja arachnoidea

Many wildflowers, including paintbrush, adorned the sandy slope as I made my arduous ascent. My tip for climbing up a scree, avoid the sandy areas and aim to climb where there are boulders instead.

Cobwebby paintbrush, Castilleja arachnoidea

Wildflowers, each in a special niche

Finally, I had made it to the ridge, and I knew I could find my way down safely.

Admiring the view from the low end of the ridge, I had to decide whether to continue to the peak or turn around. I was exhausted, it was getting late. A few hikers, father and son, greeted me as they started to descend. I asked how much farther to the summit, and they told me 45 minutes of difficult terrain. Pride urged me to keep going but I decided against it. I was dizzy, tired, and eager to get down before dark.

California tortoiseshell, Nymphalis californica

Before heading back down, I sat next to a large ice patch to fill my water bottle once more. A tortoiseshell landed directly on the ice nearby and was drinking from it. I was enthralled.

Alpine gold, Hulsea nana, the only flower I saw blooming on the ridge.

Though I didn't make the summit, it gives me motivation to return and do it again.

Lower Table Rock

noreply@blogger.com (Travis Owen) — Thu, 13 Jul 2017 04:05:00 +0000

Lower Table Rock, as seen from the road.

It's summer in Southern Oregon which means it's very hot (in the mid-90's), dry, and sunny. Working with honeybees, my current day job, is limited by the heat since temperatures above 92°F can harm the young larvae. On my way home early from work one fine, and hot, afternoon, I decided to take a more scenic route home. On a whim, I spontaneously decided to hike up Lower Table Rock, heat be damned! I've hiked up Lower Table Rock several times, all in the spring, yet I was enticed by the fact that I had never climbed up Lower Table Rock, and I had never experienced either of the Table Rocks in summer. I was determined. I was going to make this happen.

Narrow leaved milkweed (Asclepias fascicularis) and a large spider wasp (Calopompilus pyrrhomelas)

My first stop, barely twenty feet from the parking lot, was a lovely patch of narrow leaved milkweed (Asclepias fascicularis). This is the latest blooming of all the locally native milkweeds, and also has the smallest flowers. Several small pollinators were at work, including some flies and a small leaf cutter bee (Megachilidae). The most impressive and largest of the floral visitors was without a doubt a large spider wasp (Calopompilus pyrrhomelas, Pompilidae).

Narrow leaved milkweed (Asclepias fascicularis) and a large spider wasp (Calopompilus pyrrhomelas)

Spider wasps are aculeate predatory wasps which hunt living spiders for their young to feed upon as they develop. Instead of killing the spiders, pompilid wasps immobilize its spider prey by stinging it, thus rendering it paralyzed. The spider is then sealed in a chamber, normally underground, where an egg is laid and the entrance sealed. The young wasp larvae feeds on the living yet paralyzed spider until it begins to pupate. Some pompilids, such as Calopompilus pyrrhomelas, engage the spider head on in its own burrow, then turning the spiders home into its tomb. The types of spiders they hunt depends on the species of the wasp, as they are all specific.

Crown brodiaea (Brodiaea coronaria) with a large dipteran pollinator (Thevenetimyia sp.)

Further up the trail, by this point on the incline, the last of the crown brodiaeas (Brodiaea coronaria) were in bloom. Curiously, I have not observed many bees of any kind working these flowers with only a few exceptions, and have suspected flies to be important for their pollination. Low and behold, I observed several beeflies (Thevenetimyia sp.) visiting perhaps half of the brodiaea flowers. Their behavior appeared to be of the nectar gathering sort, though I do not know with certainty.

Pincushion gall (Diplolepis rosae) on a wild rose

Signs of other wasps were evident, from pompilids and ichneumonids briefly revealing themselves in the grasses, to mossy rose galls on the roses. Wild rose plants of a few select species are subject to strange growths on the stems. The growth, a gall, is formed by very small phytophagous wasps. The mossy gall wasp (Diplolepis rosae) lays an egg on or in the tissue of the new leaf buds in the spring. A reaction occurs causing the plant to form the gall, and the wasp larvae feeds on the tissue within. They overwinter inside the gall as prepupae, emerge as adults in the spring, mate, and lay eggs. Their adult lifespan is probably pretty brief, probably within a few weeks.

Royal rein orchid (Piperia transversa)

The forested incline up the side of the monument was mostly carpeted with poison oak. Every now and again I would spot tiny orchids, Piperia sp. (now considered Platanthera, but I like saying Piperia more), peeking up through the other plants. Piperia transversa leaves appear in late winter, then they usually dry up and disappear before they flower. Moths (mostly Geometridae and Noctunidae) and the occasional bumblebee is documented to visit various species of Piperia, all of which produce nectar that is held in the spur of each flower. Pollinators visit the older, lower flowers first, then move up and visit the youngest flowers at the top last. Though self pollination is possible, they set seed much better when crossed with another plant.

I was being watched

Close to the edge of the forest, which is mostly madrone (Arbutus menziesii) I heard footsteps above me. I stopped fifteen feet of a forest dweller, a black tailed deer. She just stood and stared, assessing whether I was a threat I imagine. She did,'t even flinch as I pulled out my camera to take a picture. I told her I realized this was her home, and I was her guest, and I moved along on the trail.

Western fence lizard (Sceloporus occidentalis)

Having grown up with reptiles in my house (as pets) I am always fond of seeing lizards and snakes. I grew up with green iguanas, king snakes, boas, bearded dragons, and horny toad lizards. In Southern California, we would drive out to the Anza Borrego desert where I would catch horny toads with a long piece of grass tied at the end (I'd hold them, pet them, then always let them go). Western fence lizards are very common here in Southern Oregon, so they are the most commonly encountered reptile for me. I don't catch them, except on my camera.

Lark sparrow (Chondestes grammacus)

I saw a lot of birds on the hike, but the first one I was able to photograph was the lark sparrow. This is a common bird, and I don't know much about them, so I'll leave it at that.

At last, the dry summit!

And then it happened, I came out of the forest and there I was on the summit! It was hot and dry, unsurprisingly, but a steady breeze made it feel deceptively cooler than what the actual temperature probably really was. Dry grasses and buckbrush (Ceanothus cuneatus) were the dominant plants up there, or at least the most apparent. Wyethia angustifolia and Triteleia sp. were in seed, and few other things were in flower. Clarkia amoena was practically everywhere, in bloom among the grasses, but for some odd reason I didn't take a single photo of it in bloom. The most striking part of my hike was that there were practically no other people on the trail. Maybe two other cars were in the parking lot below, and I saw only two people on the way up. Upon reaching the top, I was alone. The word serenity came to mind.

A view of Upper Table Rock, Mt. McLoughlin in the distance to the right

I diverged from the main trail and took a smaller thinner trail to the edge. Initially I was looking for more flowers, but then I realized it didn't matter and I just wanted to be on the edge and look down.

Watch that first step

Looking down, I had few words. Even now, I struggle to make sense of what I saw. There is something beautiful and awful about seeing houses, farms, neighborhoods, and businesses from a high vantage point. In one aspect, it is beautiful and awe inspiring, a scene most people (including myself) do not see very often. But for me, seeing the work of many people living their lives, seeing it from a pristine wilderness, it is sad. There is no other species on the planet which has altered their habitat so much as people. The Table Rocks might not be so special if it weren't for the great loss that has taken place over the centuries as families and entrepreneurs have expanded their uses of the land around them to suit their purposes. What surrounds the Table Rocks mostly are farms and houses, and around or within those are many agricultural weeds. The two Table Rocks are nearly pristine islands of what it all once was, surrounded by a sea of invasive sprawl. This makes me sad.

Eriogonum umbellatum

After withdrawing from the view below, I returned to focusing on what was within my literal reach. Various species of wild buckwheat, Eriogonum spp., dot the rocky edges of the Table Rocks. Most, with the exception of E. umbellatum, were in seed.

Eriogonum umbellatum with a small halictid bee

Small flies and bees were visiting the Eriogonum. Wild buckwheat has flowers with very accessible rewards, and in my experience attract a wide range of short tongued pollinators.

Western fence lizard (Sceloporus occidentalis)

Fence lizards are probably everywhere there is sun and rocks in this area. A very chubby one was pretty sure I did not see it, and allowed me to take a good close up with the sun in the background.

Common yarrow (Achillea millefolium) with a grasshopper nymph (Melanoplus sp.)

The most common insects on the summit were the grasshoppers. A beautiful green grasshopper nymph, apparently missing a leg, allowed me to photograph it on the inflorescence of yarrow. Achillea millefolium was one of the few flowers in bloom. I often see grasshoppers on flowers, it is likely they are eating flower parts rather than nectar, but one never knows. Maybe, perhaps, they are underappreciated pollinators. They are good eating for the fence lizards.

The moon, as seen from a dried vernal pool.

The vernal pools are completely dry by summer, and nearly devoid of life. Tiny fairy shrimp (Branchinecta lynchi) which once swam in these very same vernal pools earlier in the year are dead now, but their eggs are now embedded in the soil, completely immune to seasonal drought, and will come back to life in late winter or early spring when the pools are resurrected and the temperatures begin to trend upwards. For now I fathom the semblance of the moon and the dry vernal pool.

Epilobium densiflorum

Denseflower willowherb was one of the few flowers in bloom on the summit. The flowers are very ephemeral, usually only open in the mornings and then fading by noon. New flowers are produced daily.

Melanoplus devastator on tarweed (Madia sp.)

Many many grasshoppers were visible, mostly jumping away from me, perceiving me as a predator. Tarweeds (Madia spp.) were in seed, some dried to varying degrees. The grasshoppers, perhaps by coincidence, would jump away from me as I walked along the main trail, and then hit the dried stalks of a variety of plants in seed. The jostling would scatter seeds, each time grasshoppers hit a plant I could hear the tiny seeds flying everywhere.

Fitch's tarweed (Centromadia fitchii)

Resembling a starry Madia, Fitch's tarweed was mostly in seed except for a few individuals. I have seen this species growing around the base of the Table Rocks, yet seen not a single pollinator visit it.

Blister beetle (Epicauta sp.) on Fitch's tarweed (Centromadia fitchii)

In previous years I have seen blister beetles eating the flower parts of elegant tarweed (Madia elegans), and now I have seen blister beetles on Fitch's tarweed. Blister beetles are named so due to a chemical within them and on them that causes blisters when it comes into contact with human skin. Look, but do not touch.

Western mountain balm (Monardella odoratissima) above, farmland below

The main trail on the summit of Lower Table Rock is wide and straight and felt to me like it was never going to end. But it did, and I was ponce again looking down a cliff face. The rocky outcrops at the far side of the summit was host to many flowering mountain mints, Monardella odoratissima. This small mint, one of my favorite summer blooming wildflowers, found its niche in the pockets between the volcanic rocks on the edge. A few bumblebees were at work on them, but the wind and the low aspect of the sun prevented me from catching a god photo. More eriogonums were blooming on the edges, and they had me crawling along the precipice like Sméagol looking for the Precious.

Ladies tresses (Spiranthes sp.) going to seed on the summit

It was getting late, I had to turn back before it got dark. On my way back the sun was very low, and the low light highlighted the dried foliage of the grasses and other plants. I caught sight of another orchid, this time Spiranthes, ladies tresses, going to seed. These are very beautiful plants in flower, and also beautiful in seed. I hope the dust like seeds spread and grow into new plants. I've observed bumblebees visit these in flower, and unlike the other orchids in this region (Piperia, Goodyera, etc.) Spiranthes sometimes grows in open sunny fields and clearings.

Allium bolanderi, flowers withering and seeds developing

I had seen wild onions on Upper Table Rock earlier this year, Allium parvum, growing on the summit. Now in summer, those have long gone to seed, and we have the last remaining Allium bolanderi also going to seed. These grow under the canopy of the madrone forest below the summit. A smaller subspecies also grows in this region, A. bolanderi subsp. mirabile, with flowers no wider than a framing nail.

A mystery, solved

On my descent, I had seen what I thought were some kind of exotic trumpet-like flower growing under some of the buckbrush shrubs, and its identity remained a torturous mystery. Then I was informed that no, they aren't some type of amazing new flowering species that for a century has been ignored and neglected, but rather they are the beautiful seed capsules of some type of Clarkia! So I photographed the Clarkia after all! I am much relieved.

Great horned owl (Bubo virginianus)

Nearing the end of my descent, a huge owl swoops over the trail ahead of me. I was very gratified to discover this large predator had landed not fifteen feet from the trail, and posed politely for me to take a photo! Owls are amazing, and I was gifted the pleasure of encountering the largest owl in the region, the great horned owl. Nature is amazing.

Moths and Butterflies of Southwestern Oregon (2016)

noreply@blogger.com (Travis Owen) — Wed, 25 Jan 2017 15:13:00 +0000

California Sister (Adelpha californica)

When I was a young kid, before my angst-ridden teenage years, I was always excited by the sight of a butterfly, and naturally, if I saw one I attempted to capture it (unsuccessfully, however). Being mostly harmless creatures (with the exception of the stinging hairs of some caterpillars and adult moths, and a few species of moths in the genus Calyptra with piercing mouthparts), and often large and brightly colored, it is no surprise that butterflies and some moths are some of the most adored of all insects, and perhaps of all pollinators. Usually this appreciation is directed towards the butterflies (Papilionoidea), though there are also many brightly colored moths that garner our attention. In any case, butterflies are representative of one of few groups of insects that aren't arbitrarily feared.

Mossy Swallow Moth (Feralia februalis)

Moths and butterflies make up the order Lepidoptera, which literally translates to scaled wing (from the ancient Greek λεπίς ‎(lepís, “scale”) and πτερόν ‎(pterón, “wing”). This name is in view of the fact that the wings of all lepidopterans are covered in many small scales that overlap. Many have scales of various colors that together form cohesive patterns. The patterns are sometimes brightly colored, either as warnings or perhaps as a means of attracting a mate, or they are dull and help them blend in with their environment to avoid being eaten. Most but not all butterflies feed on nectar as part of their diets, as do many moths, though there are some moths which do not eat as adults (some don't even have functional mouthparts or digestive systems) and are in the adult stage only long enough to mate.

Unidentified caterpillar on Arctostaphylos viscida, a host plant to many species of Lepidoptera.

One thing most lepidopterans have in common is the larvae are plant feeders, occasionally feeding on a specific group of plants or even only on a single species. While they are in this life stage they are vulnerable to attacks from predatory vertebrates like birds or by predatory wasps and other arthropods. Parasitoid wasps often use caterpillars as hosts for their offspring. Egg(s) of parasitoid wasps are laid nearby, on, or in the caterpillar by means of an ovipositor. The wasp larvae then burrow their way through the insides of the caterpillar, eating as they go, until they are ready to exit and pupate. Though I have not seen this, many gardeners have often encountered the tomato hornworm (Manduca spp.) with dozens of small white braconid wasp cocoons on its exterior. At that point, the caterpillar is greatly weakened and will likely be unable to pupate, then die. There are also hyperparasitoid wasps which parasitize the parasitoids already within a caterpillar, possibly saving the caterpillar from being eaten alive.

An adult caddisfly, Dicosmoecus sp. (Trichoptera)

Butterflies are believed to have evolved from moths over fifty million years ago, and the first butterfly fossils are from the Eocene. Moths have been around a lot longer, the earliest fossils date to at least 190 million years old. The earliest lepidopterans are thought to have evolved from caddisflies (order Trichoptera) which have membranous wings instead of scaled wings. There are around 165,000 estimated species of Lepidoptera in the world, ~13,700 of which are true butterflies while the rest are fairly considered moths.

Edwards' Glassy-wing Moth (Pseudohemihyalea edwardsii)

There are some general differences between moths and butterflies that can be observed by the average viewer. Here I will list several differences in no order of importance. Butterflies are entirely diurnal while moths have diurnal, nocturnal, and crepuscular (active during twilight) species. The antennae are probably the most telling feature, butterflies have thin filamentous antennae with clubbed ends (sometimes hooked as is the case with the skippers) while moths typically have antennae which end in a point in the case of females or feathered in the case of some males. However, some moths (i.e. ghost moths, Hepialidae) have very short antennae that aren't readily apparent at all.

A predator of moths, I observed on one foggy fall morning as Western Bluebirds (Sialia mexicana) stalked and chased large hairy moths hiding in grassy tussocks in a field. Upon discovering the moths, the birds would pursue the moth. The moth would attempt to flee and its silhouette could be seen in the sky with several Western Bluebirds in close pursuit until its inevitable capture.

Wing color does little to help determine whether it is a moth or a butterfly, though the position of the wings at rest can be diagnostic. Most butterflies hold their wings flat together when they land while moths can either hold them to the sides, or wrapped against their bodies (as you'll see in photos below). Wing position isn't always reliably diagnostic, since there are exceptions in both moths and butterflies (i.e. Geometridae or Hesperiidae, respectively) which do not conform. Another key difference, though not readily apparent, is that while all butterflies eat as adults, many moths only eat in the larval stage and may not even have functional mouthparts or digestive systems as adults. This discounts many moths as pollinators, but they are still valuable as a food source to countless vertebrates such as songbirds and even bears.

Milkweeds are important host plants for many butterflies and moths. Asclepias cordifolia, or heartleaf milkweed, is native to California and Oregon and a host plant for monarchs (Danaus plexippus, Danainae).

Adults of many moths and caterpillars feed on nectar from flowers and are widely regarded as pollinators. However, many moths do not feed at all as adults, and of the species that do there are many other sources of nutrition that they utilize as adults. Some adults feed on honeydew or tree sap (i.e. the Mourning Cloak, Nymphalis antiopa, which prefers sap from oak trees). Many butterflies are attracted to rotting fruit and mud. Others, particularly males of both moths and butterflies, feed on fluids taken from feces or even carrion. Salts found in carrion, feces, and sweat are useful for the adult males, and usually not found in nectar. To read about creating habitat for moths and butterflies, I highly recommend the book Gardening for Butterflies: How You Can Attract and Protect Beautiful, Beneficial Insects written by the Xerxes Society (Timber Press 2016). I wrote a review for Gardening for Butterflies for the Native Plant Society of Oregon, read it here.

I have compiled all the moths and butterflies I have photographed this year and arranged them in the only way that made sense to me: taxonomically (in other words, by scientific classification). This is not all the lepidopterans in my area, merely the ones I have both seen and had the opportunity to photograph. It may come as a surprise to some that butterflies and skippers are simply a branch of this classification, merely a specialized group of moth descendants. That being said, I have decided to place butterflies at the bottom of this report. If one is highly averted to moths for some reason, they may scroll to the bottom. I, however, think moths have a lot to be appreciated, from their intricate (if occasionally dull) patterns, to their obscure lifestyles, their importance in the food web is not fully understood and they may ultimately have more to offer than butterflies. Please, enjoy.

Superfamily Alucitoidea

Family Alucitidae

The many-plumed moths are named for their wings which are composed of feather-like rigid spines from which flexible plumes radiate. This gives them a characteristic appearance. Larvae feed on honeysuckle (i.e. Lonicera hispidula) and snowberry (i.e. Symphoricarpos hesperius), both of which are present in my area in Southwestern Oregon (Rogue Valley).

Alucita sp. (January)

Alucita spp. are small moths, with a wingspan of around one centimeter. Their host plants are snowberries (Symphoricarpos spp.) which are shrubs in the honeysuckle family, Caprifoliaceae. Symphoricarpos albus var. laevigatus and Symphoricarpos rotundifolius have been documented host plants, perhaps other species as well.

Superfamily Lasiocampoidea

Family Lasiocampidae

This family is best known for the tent caterpillars, Malacosoma californicum in the western US and M. americanum in the east. Tent caterpillars build protective webs in the limbs of their tree or shrub hosts, though they aren't the only moths to do so. The fall webworms, Hyphantria cunea (Erebidae), builds similar silken nests in summer and fall while tent caterpillars make their nests in spring. In either case, the gregarious caterpillars defoliate the area within their protective webbing. Other moths in the Lasiocampidae are solitary in the larval stage.

Subfamily Macromphaliinae

Tolype distincta (September)

Tolype distincta is native to coniferous forests in the Pacific Northwest. Caterpillars feed on various conifers such as madrone (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla), and other Pinaceae, as well as having been documented on non-coniferous trees and shrubs bigpod ceanothus (Ceanothus megacarpus), mountain mahogany (Cercocarpus betuloides), and canyon live oak (Quercus chrysolepis). Adults are nocturnal and active in late summer and fall. They are drawn to lights, the individual in my photo was under a porch light the following day trying unsuccessfully to blend in with the cedar siding.

Superfamily Noctuoidea

Family Erebidae

The Erebidae, whose name has roots in Greek and Latin meaning "darkness" or "evening" reflecting when they are active, are a group of often colorful moths. This group contains some interesting members including the piercing moths (Scoliopteryginae pierce fruit to imbibe the juice, Calpinae pierce fruit for the juice as well as the skin of some mammals to feed on blood). Some members of the Erebidae have even been proven to make a sound used to deter or confuse bats, one of their primary predators. The bright colors and patterns of some species are thought to deter sight based predators such as birds, a signal that they may be toxic or distasteful.

Subfamily Arctiinae

Pseudohemihyalea edwardsii (September)

A large nocturnal moth, I found Pseudohemihyalea edwardsii clinging to a door under a porch light. The moth was nearly two inches in length, and quite conspicuous. I hope for the sake of the moth that this species is toxic since it was so highly visible. Where does this creature hide? Perhaps it is adapted to the crevices of the bark of their hosts, the oaks. Perhaps it is better hidden in the filtered light of the canopy of an oak than a blue door.

Pseudohemihyalea edwardsii has translucent wings which give it the name Edwards' Glassy-wing (September)

Pseudohemihyalea edwardsii appears to host exclusively on oak trees. Documented hosts include the species coast live oak (Quercus agrifolia), canyon live oak (Quercus chrysolepis), valley oak (Quercus lobata), and I would expect other species of oak as well within their range. P. edwardsii is found from Washington, through Oregon and California, and Arizona.

Family Noctuidae

Owlet moths, cutworms, and armyworms are the common of this large family. Owlet moth refers to the hairy bodies of some species which are reminiscent of baby owls. Cutworm or armyworm refers to the voracious and destructive appetites of some species which have sometimes been considered serious agricultural threats. Most are nocturnal and are attracted to lights, except the subfamily Agaristinae which are day flying.

Subfamily Amphipyrinae

Feralia februalis (February)

Known as mossy swallows, the genus Feralia is native to much of the United States. Feralia februalis is found from late winter to early spring in mixed coniferous woodlands in the Pacific Northwest, often in association with oaks. Their primary host plant is Oregon white oak (Quercus garryana) in Oregon, though they have also been documented on blue oak (Q. douglasii). In California they have been reported to use other plants as hosts such as California buckeye (Aesculus californica), readheart mountain lilac (Ceanothus spinosus), mountain mahogany (Cercocarpus betuloides), hollyleaf cherry (Prunus ilicifolia), and blue elderberry (Sambucus nigra ssp. caerulea). At least two other species of Feralia are native to Oregon, F. comstocki and F. deceptiva, which feed exclusively on Douglas-fir, Pseudotsuga menziesii.

Feralia februalis (February, of course!)

It is not known whether this species, or any species in the genus Feralia for that matter, feeds on nectar or feeds at all as an adult. Unlike butterflies, as previously stated, there are moths which do not feed as adults but there is a large deficiency of knowledge about which moths do or do not feed as adults.

Subfamily Noctuinae

Feltia jaculifera, attracted to lights at night, yet are active both day and night. (September)

Cutworms are considered pests by both gardeners and farmers alike. Envision a beaver gnawing away at the base of a tree and you may understand their method of feeding. Larvae are usually active on or in the soil, often emerging at night to feed and hiding during the day. Feltia jaculifera feeds on a wide variety of plant hosts, many native to its range and a few non natives of agricultural importance. The closely related F. subterranea, however, is a pest of a dozen or more plants of agricultural importance including beets, carrots, asparagus, peas, potatoes, and corn among others.

Feltia jaculifera (September)

Feltia jaculifera feeds on nectar as an adult, and may very well provide beneficial pollination services. This species has been observed feeding primarily on plants in the Asteraceae, including Joe Pye weed (Eutrochium purpureum), green-headed coneflower (Rudbeckia laciniata), nodding beggarticks (Bidens cernua), smooth beggarticks (B. laevis), and Canada goldenrod (Solidago elongata), although there are surely others of importance to this species as nectar plants. Various solitary ichneumon wasps are documented as using various species of Feltia as hosts, namely Ceratogastra ornata, Campoletis flavicincta, Enicospilus merdarius, and Spilichneumon superbus.

Subfamily Plusiinae

Autographa californica nectaring on a zinnia (October)

The genus Autographa is represented in the West by a handful of species. Some of which have seemingly limited preferences of host plants, while A. ampla and A. californica utilize many species of plants in many families as hosts. Host plants in the families Asteraceae, Fabaceae, Lamiaceae seem to be most commonly used. Trees are also often used as hosts, namely alders (Alnus spp.) and willows (Salix spp.), but also balsam poplar (Populus balsamifera), quaking aspen (P. tremuloides), birch (Betula papyrifera), chokecherry (Prunus virginiana), and viburnum (Viburnum spp.).

Autographa californica with wing damage, possibly from a bird or another predator. (October)

Autographica californica is a common species often regarded as a pest. Caterpillars of this species are known to feed on a wide variety of agriculturally important crops, though they are also abundant in woodlands and forests. Adults are mostly nocturnal, but may come out in the day to feed on nectar. The aren't picky about where their nectar comes from, and may in fact contribute to the pollination of some plants. A. californica is one of few pollinators to be documented visiting Triteleia laxa, a bulb native to California and Oregon. The braconid wasp Meteorus autographae uses Autographa spp. as hosts in the Eastern US.

Superfamily Pterophoroidea

Family Pterophoridae

The plume moths are closely related to the Alucitoidea, or many-plumed moths. Both have wings that apparently lack scales, but are rather adorned with feather-like plumes. The easiest way to differentiate between the two is how they hold their wings when they land. Many-plumed moths hold their plumes out like a fan, while pterophorid moths make a distinct "T" shape. Both are nocturnal, but may be found clinging to walls during the day. There are rumors that plume moths bite, and although it has never been substantiated there is a chance possibility that some have proboscises capable of piercing human skin (most likely male moths, but this is purely non-expert conjecture).

Pterophorid moth investigating a daffodil (March)

Plume moths visit a wide range of plants to use as hosts, the vast majority are in the Asteraceae, or sunflower family. They have also been documented using hosts in other families, such as California horkelia (Horkelia californica: Rosaceae), twinberry honeysuckle (Lonicera involucrata: Caprifoliaceae), and coyote mint (Monardella villosa: Lamiaceae). They also feed on nectar as adults. For three years I have observed pterophorids meticulously visiting the florets of Muscari spp., or grape hyacinths, in the spring at night. They have also been documented visiting yarrow (Achillea millefolium: Asteraceae), which happens to be one of their host plants.

Superfamily Sesioidea

Family Sesiidae

Sesiid moths are known as clearwing moths because most species have wings that are clear. Most are thin and somewhat small, with white, red, or yellow stripes on a black body. Many are likely wasp mimics, a strategy that may reduce predation and allows them to be active during the day. Some resemble yellowjackets (Vespula spp.) while others resemble other types of wasps. Sesiid moths in the genus Alcathoe have black bodies with orange wings, very similar in appearance to the large tarantula hawk wasps (Pepsis spp.) which share similar distributions in the American Southwest. Larvae feed within either woody plants or within roots, and some are considered serious pests.

Subfamily Sesiinae

Penstemonia clarkei (Inset: a male yellowjacket, Vespula pensylvanica)

Penstemonia clarkei is a small moth which, as the generic epithet suggests, seeks hosts in the plant genus Penstemon or the related Keckiella. Penstemonia larvae feed within the crown of their hosts, often to the detriment of the host. Penstemonia spp. are wasp mimics (compare to the inset image of a male Western yellowjacket, Vespula pensylvanica).

Superfamily Geometroidea

Family Geometridae

Geometrid moths are so named for the mode of locomotion of their caterpillars, also known as inchworms or loopers. "Geometridae" derives from the Greek word γεωμέτρης (geometer, or earth-measurer). Most caterpillars feed on plant foliage, and adults are typically but not always nocturnal. Adults are attracted to lights.

Subfamily Ennominae

Cochisea sinuaria (September)

Although dull in color, Cochisea sinuaria is rich in pattern. I found this specimen clung to the wall of an outbuilding under a light, but I imagine against the bark of a tree it would be nearly indistinguishable. C. sinuaria larvae feed primarily on the leaves of manzanita, particularly Arctostaphylos viscida and A. glauca. The species also has been associated with other potential host plants deerbrush (Ceanothus integerrimus), coastal sage scrub oak (Quercus dumosa), mountain mahogany (Cercocarpus betuloides), and laurel sumac (Malosma laurina).

Tetracis jubararia (October)

Tetracis jubararia is a twig mimic as a caterpillar. Adults are active in late summer and fall, when they lay eggs and perish. Eggs endure winter, and hatch the following year. T. jubararia has many hosts, usually broadleaf trees, but also many shrubs. A few of the host genera include maples (Acer spp.), alders (Alnus spp.), birches (Betula spp.), dogwood (Cornus spp.), cottonwood (Populus spp.), currants (Ribes spp.), azaleas (Rhododendron spp.), willows (Salix spp.), elderberry (Sambucus spp.), snowberry (Symphoricarpos spp.), and huckleberries (Vaccinium spp.) among many others.

Subfamily Geometrinae

Nemoria darwiniata (September)

The small Nemoria darwiniata are common in wet or dry woodlands in much of North America. I found this one in an apiary while working some hives, hence the sugar-coated glove (the moth was on my finger).

Nemoria darwiniata (September)

Nemoria darwiniata has many hosts, usually broadleaf trees and shrubs. Hosts include madrone (Arbutus menziesii), manzanita (Arctostaphylos spp.), wild lilac (Ceanothus spp.), golden chinquapin (Chrysolepis chrysophylla), yellow rabbitbrush (Chrysothamnus viscidiflorus), oceanspray (Holodiscus discolor), California wax myrtle (Morella californica), bitterbrush (Purshia tridentata), oaks (Quercus spp.), and willows (Salix spp.) As the caterpillar feeds, it takes on the color of the host plant, which is common for many moths as a defensive strategy.

Subfamily Larentiinae

Costaconvexa centrostrigaria (October)

The Bent-line carpet moth, Costaconvexa centrostrigaria, is a small American species, in fact the only species of Costaconvexa in the US, although it is widespread. Adults fly from May to October in the Northwest, but all year in the southern distribution of the species. Adults are also drawn to light, which is where I found this individual.

Costaconvexa centrostrigaria (October)

Larvae of Costaconvexa centrostrigaria feed exclusively on Polygonum species (knotweed) and also the closely related Persicaria hydropiper (formerly included in the genus Polygonum).

Superfamily Pyraloidea

Family Pyralidae

The family Pyralidae has some of the most diversity in life histories of all lepidopterans. Diets vary widely, from phytophagous species to species that live parasitically in ant nests (Wurthiinae spp.), predatory species (certain Phycitinae spp. prey on scale insects), or live in the nests of bees as inquilines or kleptoparasites. Some species spend their larval stage under water (Acentropinae) while others are adapted to very dry environments, such as living off of dried stored food products (some Phycitinae and Pyralinae) and are considered to be pests.

Subfamily Galleriinae

Galleria mellonella on wax comb of Apis mellifera

Galleria mellonella is better known as the greater wax moth, most often regarded as a pest of honey bee colonies. They are one of a handful of moths which live within apid bee nests, that is bees within the family Apidae which build wax comb (i.e. bumblebees, honeybees). Wax moths are also occasionally found within old vespid wasp nests. Both the greater and lesser wax moths (Galleria mellonella and Achroia grisella, respectively) are pests within honeybee colonies, as well as in the nests of stingless bees (Meliponini). Aphomia sociella is a pest of bumblebee nests in the UK and perhaps the Eastern US as an introduced species, only rarely a pest in honey bee colonies.

Galleria mellonella

The greater wax moth is not all bad, though the species does create a headache for beekeepers. The larvae serve as hosts for braconid and ichneumon wasps (Habrobracon hebetor and Pimpla turionellae, respectively, and perhaps others). Vespid wasps, such as yellowjackets and paper wasps (i.e. Vespula and Polistes spp., respectively) prey on the larvae of wax moths. The greater wax moth is also used in laboratories as infection models for the study of bacterial diseases and for antimicrobial drug testing. In this way they replace vertebrates, particularly rodents, which have presented many ethical issues as well as time and budgetary constraints. Few ethical restrictions keep researchers from running tests on wax moth larvae, and this may even incite some excitement from beekeepers who despise the moths.

Galleria mellonella larval silk and frass on honeybee comb

Greater wax moth females may lay over a thousand eggs in their brief adulthood, around twelve days. Adults are nocturnal, and probably attracted to the scent of beeswax. Many dozens of larvae may be found within a single frame tunneling below the surface of the comb. They build a protective layer of silk between frames, perhaps to allow easy passage from one comb to the next. The entire silken structure is laden with frass, or feces. In the wild the moths serve an interesting ecological role. When a wild or feral honey bee colony dies, the moths move in and destroy everything, leaving a nice clean cavity for new bees to move into rather than using the old comb which may be tainted with disease.

Galleria mellonella larval silk and frass

Typically the best defence against wax moths is to keep strong colonies. Populous and healthy honey bee hives will fight off any intruder. Stored comb is the most susceptible to wax moths. Cold temperatures are a good preventative measure as the moths require warmth to mate and lay eggs. Immature moths go through winter as either prepupae or perhaps as eggs if they are in protected locations. That said, certain strains of BT (Bacillus thuringiensis) are effective against wax moths.

Superfamily Papilionoidea

The superfamily Papilionoidea contains all of the butterflies, yet an alternative taxonomic treatment places the skippers (Hesperiidae) in a separate superfamily, Hesperioidea. Butterflies differ from the moths by a few key characteristics, and a number of other variable characteristics which can be used to support the key differences. Butterflies, unlike moths, have clubbed or hooked antennae, in other words the antennae end thicker than they began. Moths have antennae that taper to a point or are feathered in some males. Another key difference, moths have a frenulum, though not readily visible unless a moth is in hand, is a wing-coupling device that ensures the hind and forewings work in unison. Butterflies lack this feature. Other differences are variable, though may be accounted for together to make a positive identification. Butterflies are diurnal (occasionally crepuscular), yet moths are active both day and night depending on the species. Most butterflies are colorful, and most moths are dull colored. However, there are many drab butterflies and many colorful moths (particularly tropical moths). In the pupal stage, moths and butterflies differ in that moths tend to build silken cocoons while butterflies tend to build a chrysalis, often smooth with a hardened surface. This, of course, is variable as some moths also have pupal cases that are smooth and shell-like.

Family Lycaenidae

The family Lycaenidae, which means "little butterflies" due to the small size of many species, are widespread yet contain many rare and uncommon species. Known as the hairstreaks, blues, and coppers, they often have delicate wing markings. While many species are phytophagous, there are also predatory and parasitic species. Many species have mutualistic relationships with ants, where in exchange for protection by the ants the caterpillar secretes a complex sugary substance. Some of these caterpillars move directly into the ant mound where they continue to excrete the sugary substance sought by the ants. These caterpillars within the ants nest often become parasitoids and feed on the ants within the nest! Other lycaenids switch from a plant based diet to a predatory diet of aphids being tended by ants. Lycaenids often pupate underground, often within an ant colony, and they must emerge from the mound to fully expand and dry their new wings once reaching adulthood.

Subfamily Theclinae

Strymon melinus (June)

The gray hairstreak is a very common and widespread butterfly, and one of the most encountered of the hairstreaks. Their abundance may be due in part to their adaptability. They have a ridiculously wide range of host plants, and can be found nectaring on a similarly large array of plants. The caterpillars tend to take on the color of the host plant. They are occasionally considered pests as they have a preference for legumes, many agriculturally important. They are active from spring to fall, found in disturbed habitats, woodlands, and agriculturally dominated areas.

Mating flower longhorn beetles (Strophiona tigrina, upper left) and Strymon melinus (lower right)

Hosts vary widely from natives to nonnative plants, including garden plants and noxious weeds. In some areas they have even been shown to be dependant on exotic plants where native plants are uncommon, such as urban and suburban areas. Thus, an attempt to eradicate a neighborhood of all naturalized weeds may have a negative effect on the butterfly population. This suggests the need for programs to reintroduce native plants into urban and suburban areas, which will support a greater diversity of native pollinators.

Strymon melinus (upper left) and Bombus vandykei (lower right) on Allium ampeloprasum var. ampeloprasum (July)

Gray hairstreaks aren't picky about nectar, and will visit a wide array of plants for it. I personally have observed them visiting ornamental onions (Allium spp.) and wild buckwheat (Eriogonum spp.), though they have been documented visiting a wide range of flowering plants, many with easily accessible nectar.

Family Hesperiidae

The skippers, occasionally placed in their own superfamily (Hesperioidea), are typically small and short-bodied, hairy butterflies with a rapid skipping flight pattern. Many skippers are understandably confused for moths, their compact and hairy forms coupled with the position many of the skippers hold their wings. Many skippers hold their wings differently than most butterflies, instead of holding them flat together or in a flat plane many skippers hold their wings in a "V" shape. Many skippers are also comparatively plain in color, yet they still have the clubbed antennae similar to other butterflies, except skippers have a little pointed hook at the tip as well. Many skippers lay their eggs by dropping them on/near their host plant(s) one egg at a time, doing many passes to lay a sufficient number of eggs.

Subfamily Hesperiinae

Ochlodes sylvanoides (September)

The woodland skipper is a grass skipper native to the Western US as far east as North Dakota and north from British Columbia to Saskatchewan. They are primarily active in late summer and fall, during which time they imbibe nectar, find mates, and lay eggs, all in a matter of weeks. First instar caterpillars hibernate through winter, complete their feeding in the following spring, go through summer in diapause, then build their chrysalis and pupate to emerge as adults in the fall. They seek nectar from a variety of plants, and males occasionally feed on carrion.

Ochlodes sylvanoides (September)

Grass skippers (Hesperiinae) use host plants in the grass family (Poaceae). The woodland skipper uses a variety of grasses and related plants as hosts, from Bermuda grass (Cynodon dactylon), canary grass (Phalaris spp., including the invasive P. arundinacea, but also Western natives P. californica and P. lemmoni), Colonial bentgrass (Agrostis capillaris), Johnson grass (Sorghum halepense), wildrye (Elymus cinereus, E. condensatus, E. trachycaulus ssp. trachycaulus, and E. triticoides), and wheatgrass (Agropyron sp.). The fact that some of these species are nonnative, even noxious, in Oregon and California speaks to the adaptivity of some butterflies due to their ability to utilize hosts they didn't coevolve with.

Ochlodes sylvanoides (September)

In 2016 I observed many woodland skippers nectaring on a pungent mint-relative, Trichostemma lanceolatum also known as vinegar weed (though the smell is more akin to gasolene). The plants are traditionally considered to be bee pollinated, yet I observed pollen being deposited consistently onto the top of the thorax of the skippers. Pollination seemed a likely scenario, though I can not confirm this with seed viability (I did not check).

Hesperia juba (October)

Another common skipper, the juba skipper is a relatively large skipper with a green body. They are typically a montane species but will move to lower altitudes. Females lay eggs singly, and caterpillars fold leaves together with silk to protect themselves. Caterpillars probably hibernate through winter, and may even develop under snow. They visit a huge variety of plants for nectar, the most frequently observed skipper in my area. Known hosts include bluegrass (Poa pratensis et al), bromes (Bromus inermis, B. rubens, and B. carinatus), Nevada needlegrass (Stipa nevadensis), reed canary grass (Phalaris arundinacea), slender hairgrass (Deschampsia elongata), and Tridens spp.

Family Pieridae

The whites, sulphers, and yellows comprise the family Pieridae. Unsurprisingly, they come in the colors white, yellow, orange, and occasionally black and many have ultraviolet markings used in courtship. They are mostly native to the tropics though there around sixty species native to North America. Some are introduced pests of plants in the cabbage family (cabbage, mustard, etc.) and legumes.

Subfamily Pierinae

Pieris rapae (October)

The cabbage white, Pieris rapae, is an introduced species. Caterpillars are very common pests of mustard family plants in spring and summer. Adults are common from spring to fall, widely distributed throughout the US but mostly found in agricultural and urban areas. They seek nectar on a variety of plants, both native and exotic. Caterpillars of P. rapea and the related P. brassicae are parasitized by the braconid wasp Cotesia glomerata.

Family Nymphalidae

The brush-footed butterflies are so named because their front two legs are much reduced in size and have small brushes that may be used to enhance their ability to smell or taste. In appearance they have only four legs because they use only four legs to walk, the two front legs are kept tight against their bodies when not in use. While many feed on flower nectar, many other feed mostly or solely on tree sap, rotting fruit, dung, or carrion. Brushfoots overwinter as adults or sometimes as larvae.

Subfamily Limenitidinae

Adelpha californica (syn. A. bredowii) taking up sweat from my finger (October)

The California Sister (Adelpha californica) is found in montane forests and oak woodlands from Western Oregon to Southern California. Caterpillars are common in late spring, when they feed on oaks (Quercus spp.), tanoak (Notholithocarpus densiflorus), and western chinquapin (Chrysolepis chrysophylla), all of which in the Fagaceae.

Adelpha californica (October)

The California Sister feeds mostly on honeydew and tree sap, though they occasionally take nectar from milkweeds, plants in the Asteraceae, or a variety of others. On the day I photographed this butterfly, it was attracted to the top bars of the frames in an empty beehive. The sweet smell of the propolis or other honeybee residues may have offered the butterfly a foodsource. Propolis, after all, is made from tree sap and plant resins.

Adelpha californica (October)

With some patience, and by moving very slowly, I had coaxed this butterfly to climb onto my finger. There it remained for over ten minutes, continuously probing my finger with its propolis. I had been working that day, so it may have been lapping up sweat. Aside from sap and honeydew, the California Sister will occasionally feed on sweat or animal scat, a source of salts and other minerals not found in sap, nectar, or honeydew.

Subfamily Nymphalinae

Junonia coenia with a damaged wing (September)

The common buckeye is common in most of the United States and eastern Canada, but in the West it is only found as far north as Oregon. They use a variety of host plants in a handful of families. Most of their host plants are in the Plantaginaceae (Antirrhinum, Gambelia speciosa, Hippuris vulgaris, Maurandya antirrhiniflora, Penstemon, Plantago, Veronica, and Kickxia spp.) and Scrophulariaceae (Cordylanthus rigidus subsp. rigidus, Keckiella, Nuttallanthus texanus, Scrophularia californica, and Linaria spp.), though they have also been known to utilize paintbrush (Castilleja spp., Orobanchaceae), monkeyflower (Mimulus spp., Phrymaceae) and western vervain (Verbena lasiostachys, Verbenaceae). They aren't strong fliers, but I had observed on more than one occasion that the common butterfly was more skittish than others.

Junonia coenia (October)

Peculiarly, common buckeyes have been documented by many observers as visiting various grasses and probing them with their proboscis as if they are attaining something from them. Poaceae do not produce nectar, being purely wind pollinated, so it is a mystery. The grasses yellow bluestem (Bothriochloa ischaemum var. songarica), spear grass (Heteropogon contortus), and dallisgrass (Paspalum dilatatum) have been documented as species visited by the buckeye. Two possibilities seem plausible to me, the first being that there are sap sucking insects leaving behind honeydew, and the second being that the grass sap had been exposed somehow (like a broken stem). Yet, a mystery it remains.

Wasps of Southern Oregon (2016)

noreply@blogger.com (Travis Owen) — Sun, 01 Jan 2017 17:19:00 +0000

Mischocyttarus flavitarsis ♀ eating honey.

At first mention of wasps, many people withdraw to a position of fear. Wasps can be intimidating to some, notably the notorious (infamous?) social wasps in the Vespidae (yellowjackets, hornets, and paper wasps). Yet, the vespid wasps only represent around five percent of total wasp species. The majority of the world's wasps remain unseen, some are too small to see with the naked eye (i.e. Mymaridae) while others are large enough to take down a tarantula (Pepsinae). Two of smallest flying insects in the world are parasitoid wasps in the genus Megaphragma (Trichogrammatidae) which are smaller than some amoebae, single celled organisms, and are parasitoids of the eggs of thrips which are themselves incredibly tiny (the smallest arthropods are 139 µm).

Polistes aurifer, a docile paper wasp which forms small nests

Wasps are incredibly diverse and fill many ecological roles. They are pollinators, predators, parasitoids, and also food for vertebrates including birds and reptiles (as well as feeding on each other, in some cases). Many of these attributes are overlooked, the deeds of a few of the eusocial stinging predatory wasps has a deep impact on the psyche of many people. The fear induced by the eusocial wasps is directly tied to their survival, the nests of paper wasps and similar colony forming wasps are full of precious brood and occasionally other resources that attract a myriad of predators, including but not limited to bears and birds. The venom produced by the eusocial wasps differs from the solitary wasps in its mode of action. The solitary wasps, even the very painful sting of the tarantula hawks (Pepsis spp.), has little to no lasting symptoms or damage after the pain subsides. Eusocial wasps, however, have venom that breaks down cell membranes and damages nerves, and with enough stings, can stop a heart from beating and kill. Not all wasps have stingers, while males of all hymenopterans lack stingers.

Ophion sp. (Ichneumonidae: Parasitica), a parasitoid wasp that uses caterpillars as hosts for its larvae.

The Hymenoptera is broken into three groups: Symphyta, Parasitica, and Aculeata. The first and most ancient of these is the Symphyta which contains the sawflies, horntails, and wood wasps (not pictured). It is believed that all other hymenopteran groups descended from this clade. The Symphyta do not have stingers, but an ovipositor used to bore holes and lay eggs in plant material (it is saw-like, thus sawfly). The Parasitica is comprised of true wasps (which have the characteristic constricted wasp waist) that do not have true stingers but ovipositors instead. Many are parasitoids or predatory in the larval stage, and all are solitary (do not form colonies). The last group, Aculeata, contains the most familiar and advanced members of the Hymenoptera: stinging (predatory) wasps, bees, and ants. Most have stingers but lack ovipositors, eggs are dropped out of a special opening at the tip of the abdomen under the stinger. One exception is the Chrysidoidea which, although belonging to the Aculeata, generally lack or have greatly reduced stingers and venom glands.

Philanthus crabroniformis ♀ flying towards Eriogonum compositum

As pollinators, wasps are highly underappreciated. Wasps visit many flowers in many plant families for nectar, their primary source of carbohydrates as adults. A few species also collect pollen though pollen collecting is mostly limited to pollen wasps (Masarinae, more like bees in their dietary habits than wasps). Most wasps prefer flowers with easily accessible floral rewards, such as Eriogonum, Ericameria, Daucus, and Levisticum among many others. Australian orchids in the genus Drakaea are an example of plants that are strictly pollinated by wasps, specifically in the family Thynnidae (Vespoidea), by means of mimicry. Drakaea flowers resemble the flightless female wasps which the males attempt to carry to a secluded location to mate. Through several frustrating attempts, Drakaea orchids are pollinated and the duped males receive nothing for their efforts.

Eumenes verticalis, a solitary potter wasp that builds small urn-shaped nests to lay its eggs in.

I have observed a variety of plants that are visited by wasps. Most have easily accessible nectar as most wasps have relatively short proboscises. The majority of the wasps I've observed on flowers are predatory wasps (Aculeata) although parasitic wasps (Parasitica) are also known to visit flowers but are less frequently seen. The few parasitic wasps I've observed on flowers were an American pelecinid wasp (Pelecinus polyturator: Proctotrupoidea) on Daucus carota and a carrot wasp (Gasteruption sp.: Evanioidea), ironically, on Allium cepa. There are orchids which are primarily pollinated by parasitoid wasps, such as the Cryptostylis orchids whose only known pollinators are male wasps of the genus Lissopimpla (Ichneumonidae). Some wasps don't feed on nectar, instead some feed on honeydew (the frass of sap-sucking insects) or on the hemolymph (arthropod equivalent to blood) of hosts or prey. A few wasps (some Cynipoidea and Trichogrammatidae) don't feed at all in the adult stage, and only live long enough to mate and lay eggs.

The big hairy bee-hind of Bombus vosnesenskii (Apinae) foraging on Lathyrus latifolius. Branched hairs are what differentiate bees from wasps, a feature bees adapted to collect pollen as the primary source of protein for their larvae. Wasps are carnivorous in the larval stage (with the exception of pollen wasps and plant tissue feeders like the cynipid wasps).

Bees likely diverged from sphecid wasps in the early Cretaceous period (see Bees of Southern Oregon, Owen 2016), evolving branched hairs for collecting pollen. Wasps are traditionally carnivorous, and retained simple unbranched hairs. Thus, wasps are not as efficient as pollinators when compared to bees. However, insect-pollinated angiosperms have adapted pollen that is often slightly sticky which helps it adhere loosely to insect bodies. Additionally, flying insects collect a strong positive charge from flight, while plants (which are grounded, literally) have a weakly negative charge. When the positively charged flying insect contacts the negatively charged flower, pollen adheres to the insect with static electricity and the flower temporarily picks up a positive charge. This change of electric charge can also be detected by some pollinators, such as bees, who will avoid the positively charged flower until the negative charge is restored. A single grain of pollen placed on a receptive stigma is all that is required to produce a fertile seed, so efficiency may not be as important as frequency of flower visitation and flower constancy (when a pollinator forages on a single plant species rather than many different species).

Polistes aurifer nest, docile as can be.

It is unfortunate that wasps are so quickly dismissed as pests. Ironically, wasps contribute in a big way to our pest control efforts by preying on many other arthropods that are considered true pests. Predatory wasps are predators of the larvae of many insects from cutworms, to caterpillars, to wood boring beetle larvae and non insects like spiders. Parasitoid wasps, largely unseen, use hosts from nearly every insect order as well as non-insect arthropods, hosts such as mites, aphids, cockroaches, dragonflies, grasshoppers, thrips, barklice, and antlions to name a few. The pest control services of all insects in the United States is estimated to be around $4.5 billion considering the losses due to pests if the current services were ceased (Losey & Vaughan 2006). The Hymenoptera account for at least sixty percent of all insect predation on other insects (LaSalle & Gauld 1991), so it could be extrapolated that around $2.7 billion of annual pest control services can be attributed to the Hymenoptera.

Unidentified ichneumon wasp (October)

Below I have composed all the wasps I have been fortunate enough to photograph and identify this year in my Southwestern Oregon locale. They are arranged by one of several currently accepted (or nearly so) taxonomic interpretations. A fair warning, this is not an at all a comprehensive account of the wasps in my area, only the ones I have observed and photographed. The first section, Aculeata, is composed of the most familiar of the wasps: the stinging (or predatory) wasps which include the social wasps and also the bees (I have treated bees in a separate composition, see Bees of 2016). The following section, Parasitica, contains the parasitoid wasps which do not have stingers as the predatory wasps (Aculeata), although some are capable of poking and injecting venom. Lastly I have included ants, though technically they are considered to be a family within the Aculeata. Information of their life histories has been compiled from a variety of sources, found in the References at the end. I have also included many personal observations and speculation, if nothing else, for my own amusement. Please, enjoy:

Aculeata

Eumenes verticalis ♀

The Aculeata represents all the stinging Hymenoptera, the females of which have stingers but lack ovipositors, unlike the Parasitica which is comprised of parasitoid wasps which lack stingers but have functional ovipositors. Some species (many Chrysidoidea) have lost their stingers and evolved unique forms of ovipositors out of their abdominal segments rather than from the type of ovipositor found in the Parasitica (which evolved into the stingers of the Aculeata). Many in the Aculeata can fairly be called parasitoids or kleptoparasitoids (i.e. Chrysidoidea, Ampulicidae, etc.) as many use living hosts to raise their young unlike the typical predatory wasps in most genera. Eusociality, complete with divisions of labor and occasionally nonreproductive individuals such as what is found in honeybee colonies, is found exclusively in the Aculeata. This is the most advanced of the clades of the Hymenoptera, and far more is known about their biology than the of the many obscure wasps of the Parasitica.

Superfamily Apoidea

The Apoidea contains the bees (Anthophila) and many of the bees closest relatives: Ampulicidae, Sphecidae and Crabronidae. These non-bee groups contain predatory wasps such as the sand wasps (Bembicini) and mud daubers (Sceliphrini). Wasps capture prey from various insect orders to bring back to the nest, including cockroach nymphs, aphids, beetles, butterflies, and even bees and other wasps, as well as other arthropods like spiders. Some are kleptoparasitic and will steal prey from the nests of other wasps. Many nest in the ground, but still vary widely in their construction. Cockroach wasps (Ampulicidae) paralyze cockroach nymphs to drag back to the nest, much like pomplid wasps drag spiders, where an egg will be laid on it and the cell sealed. Once hatched, the larvae will feed on the still living, but paralyzed, roach nymph until it pupates.

Family Crabronidae

The Crabronidae is probably the most closely related family to the Apidae (bees), though it is entirely predatory. This group differs from the Sphecidae, which it was once included, most notably by the lack of a stalked petiole, or thread-waist found in many wasps, between the thorax and abdomen. Many are bee-like in appearance due to the short petiole, but have very little hair on their bodies and so appear more shiny than most bees.

Subfamily Philanthinae

The subfamily Philanthinae are predators of bees (in the tribe Philanthini), ants (tribe Aphilanthopini), or beetles (tribe Cercerini). They are often seen at flowers where adults drink nectar, and seek prey for their nests in some instances. There are many species in this group, one of the largest in the Crabronidae, and they all nest in the ground. They are all solitary, though they occasionally nest gregariously.

While the tribe Philanthini contains bee predators, they occasionally collect other wasps to provision their nests. The tribe Cercerini is particularly fond of weevils. The tribe Aphilanthopini are specialists who prey on ants. The genus Aphilanthops (Aphilanthopini) provision nests exclusively with winged queen ants in genus Formica. Often many prey insects are taken per cell, highly dependant on the size of the prey. For larger prey, less are needed.

Tribe Philanthini

The tribe Philanthini contains three genera: Philanthus, Prophilanthus, and Trachypus, although Philanthus is the most studied of these. The genus Philanthus contains about 135 species in the world with thirty two of them native to the United States and Canada. They dig one to several nests in sandy or hard packed alluvial soil each season, often over a period of a few weeks to a month, depending on the species. Each female excavates and provisions her own nest, though many nests may be found very close together where the substrate is suitable. A nest may have between four to fifteen separate cell chambers which branch off of a main tunnel. When a female completes and provisions one nest, she may go on to build more nests if she lives long enough.

Philanthus crabroniformis ♀ and a carpet beetle (Dermestidae) on Eriogonum compositum (June)

Philanthus spp. are known as beewolves, a common name which depicts their prey of choice. The types of bees hunted vary regionally and between species, although bees in the family Halictidae are most common, but other bees in the Colletidae, Andrenidae, Megachilidae, Anthophoridae, and Apidae are also occasionally hunted. Honeybees are more typical prey of larger Philanthus species such as P. bicinctus, P. crabroniformis, and P. sanbomii, as well as the European P. triangulum. Other wasps are occasionally captured and subdued as prey to be placed in a cell. Wasps in the families Scelionidae, Eumenidae, Masaridae, Sphecidae, Ichneumonidae, Braconidae, and Chrysididae have been found in the nests of various species of Philanthus.

Philanthus crabroniformis ♀ on Eriogonum compositum

Beewolves pursue bees (or other wasps) which are in flight more often than when the prey are visiting flowers, although flower patches are the usual hunting grounds. Philanthus are heavily armored and resistant to stings from bees but are still occasionally killed in action. Prey are often stung quickly resulting in complete paralysis, ensuring "fresh meat" for the developing Philanthus larvae. Between six and twenty four prey items (individual bees or wasp prey, that is) are placed into each cell. The amount of prey items used in each cell probably depends on the size of both hunter and hunted. Once the last bee is placed in the cell, an egg is laid and the cell is sealed with a soil plug.

Philanthus crabroniformis ♀ on Eriogonum compositum

Male beewolves (not pictured) have large eyes which nearly touch at the top of their heads, much like the eyes of male honeybees (Apis mellifera ♂, also known as drones). Males are territorial, and mark their territory with volatile chemicals secreted from their head. The scent is spread onto the abdomen with specialized brushes on the head, and distributed on plant material by dragging the abdomen. When a trespasser is detected, a male will meet its opponent with head butting and grappling. Since they can't sting, males of some species will bite other males who enter their territory, although Philanthus crabroniformis isn't known to do so.

Philanthus crabroniformis on Croton setigerus (July)

I have encountered Philanthus crabroniformis twice, both times visiting flowers. Eriogonum compositum (Polygonaceae) is a relatively large clump forming perennial that grows in rocky places near waterways in my region (Southern Oregon). E. compositum has a generalist pollination strategy, attracting a wide range of pollinators with the exception of bees which were scarce during my observations despite being located near a large honeybee apiary. Croton setigerus (Euphorbiaceae) is a low growing annual herb which grows in fields and roadsides, often places with little competition and poor conditions. C. setigerus, too, appeared to have a generalist pollination strategy, although it attracted fewer insects than E. compositum. Halictid bees (Halictus ligatus) and the occasional honeybee were observed visiting the male flowers, while the odd ammophiline wasp was observed along with P. crabroniformis.

Family Sphecidae

Members of the family Sphecidae are commonly known as thread-waisted or sand wasps, delineating their morphology and nest biology. Contrary to the Crabronidae, sphecid wasps have a thin and elongated petiole (the waist). Most species nest in the ground in sparsely vegetated areas, but a few build mud nests above ground (the aptly-named mud daubers). A few species nest in hollow plant stems or abandoned bee tunnels in wood. They are solitary, and similar to the Crabronidae many mass provision nests with paralyzed arthropods (larval and adult) from various taxonomic orders including spiders (Araneae), grasshoppers (Orthoptera), beetle larvae (Coleoptera), and caterpillars or sawfly larvae (Lepidoptera or Hymenoptera) depending on the wasp species. Adult wasps feed on flower nectar, and will also feed on honeydew, extrafloral nectaries, and the hemolymph (arthropod blood) of their hosts. Some species place a single large prey item in a cell chamber, but the number of prey items depends on the size of the wasp and the size of the prey.

Subfamily Ammophilinae

The Ammophilinae are ground nesting wasps most commonly referred to as the thread-waisted wasps, although other families share this trait. Females excavate unbranched tunnels in sandy compacted soil with one to two cell chambers. Nests are provisioned with one large to many small moth caterpillars or sawfly larvae (similar in appearance to lepidopteran caterpillars) either all at once or progressively as the wasp larvae grows.

Ammophila sp. on Allium sphaerocephalon (June)

Ammophila spp. excavate nests with the use of their mandibles and legs, often taking the excavated soil away from the nest entrance. A female will first dig a nest then use a pebble and sand to conceal the entrance before embarking on the task of finding a caterpillar host. Once paralyzing a host and returning to the nest, she will reopen the nest and drag the caterpillar inside. Once enough hosts are delivered into the nest, or a single large caterpillar host is delivered, the female will lay an egg and seal the nest again with a pebble and cover it in sand to hide it. If the host caterpillar is large, the female wasp will be unable to fly with it and must drag the prey back to the nest, occasionally at incredible distances from the nest site. How they can find their way back to the nest whilst dragging their prey is a mystery. It would be similar to taking a plane from a house you just moved into and then walking backwards back home.

Ammophila sp. on Allium sphaerocephalon (June)

Adult Ammophila wasps feed on flower nectar, and so are often found visiting flowers. I have seen them on a variety of flowers including various alliums (Allium spp.), Croton setigerus, and various mint relatives (e.g. Origanum spp.). I would assume they have excellent eyesight as they have made themselves very difficult for me to get a satisfactory photograph.

Subfamily Sceliphrinae

Nearly 150 species in six genera around the world and seven species in three genera north of Mexico make up the subfamily Sceliphrinae. Wasps in this subfamily live in a diversity of nest types including exposed mud nests (Sceliphron spp.) or preexisting holes in wood (Chalybion spp.). Prey are paralysed with one or more stings then sealed in the nest for the larvae to feed on. One to twenty or more prey items are placed in cells depending on the size of the prey. Prey includes spiders or cockroaches (Podium spp.).

Tribe Sceliphrini

Sceliphron caementarium (August)

The black and yellow mud dauber (Sceliphron caementarium) is a common species of solitary wasp. They are large wasps, over an inch long (~28mm) with a thin and highly elongated petiole. Though they may appear daunting, stings by mud daubers is uncommon. The sting is usually reserved for subduing and paralyzing hosts for their offspring: spiders. Spiders in many families have been found in mud dauber nests including the families Anyphaenidae, Oxyopidae, Salticidae, and Thomisidae. They have a marked preference for orb weavers, Araneidae, whose stationary lifestyle may facilitate a relatively easy capture. The female wasp lays an egg on the first spider to be placed in a cell, then continues to place up to twenty five or more spiders inside until the cell is full. The wasp larva begins feeding on the organs and other internal structures of the spiders before eating the rest of the body. The spider provisions are consumed within two to three weeks. I've often found a few dessicated spider remains inside the cells, even after the wasps have built their pupal case.

Sceliphron caementarium nest is composed of layers of mud.

The nests of S. caementarium are the most conspicuous sign of their presence, being made of locally sourced mud and adhered to a variety of locations, some of which highly visible. Nests start as a single tube-like cell, then more cells are added. Sceliphron requires a source of mud nearby, in contrast to related genera that bring water to mineral soil to make their own mud for nest building. Once all the cells are provisioned and sealed, the mother wasp covers the entire structure in a thick second layer of mud. The mud served to protect the larvae from parasites and predators, as well as to regulate the conditions inside (protect from frost, etc.).

Sceliphron caementarium nest variation

There is great variation among nest shape and sizes. Some nests I've found are just a single capped cell while others may be several inches long and have two dozen cells. The type of mud available determines the color of the nest, and to an extent the shape and texture as different forms of mud have different working properties.

Sceliphron caementarium pupal case with prepupae within

There may be one to a few generations per year, but the last generation appears to enter a state of diapause in the prepupal state after building the pupal case. The case, or puparium, is spun from a fine silk-like material and forms a soft cellophane-like shell that protects the prepupae from excessive moisture and pests. The case itself is suspended within the mud cell, sometimes with the dessicated remains of a few spiders.

Contents of Sceliphron caementarium nest with prepupae (within and without puparium, left) and a cuckoo wasp (Chrysididae) adult and prepupal case (right).

Nests of Sceliphron caementarium are often targeted by parasites or are subject to damage from microbial life such as mold or fungi which seem to destroy the viability of individual cells. Interestingly, within a single nest, some of the individual cells may be subjected to various maladies while others may remain healthy and viable. When dissecting a nest in early winter, I discovered a green metallic cuckoo wasp (Chrysididae, see below) dead in one of the cells. Several of the cells also had an immature chrysidid prepupa in a light cream-colored pupal casing instead of the elongated amber-colored pupal casing of Sceliphron.

Superfamily Chrysidoidea

The Chrysidoidea, often called cuckoo wasps, are a unique group of parasitoids and kleptoparasites within the Aculeata. The lives of some are nearly identical to the parasitoid wasps in the Parasitica though they differ in that they had developed stingers (like all aculeate wasps) from the parasitican ovipositor. Some, however, lack functional stingers (or have stingers that are much reduced) and have highly reduced venom reservoirs. Obversely, some chrysidoids (i.e. Bethylidae) are said to have a painful sting while others have enlarged venom glands.

While some, such as the Chrysididae, are kleptoparasites which lay their eggs in the nests of other bees or wasps, the rest could fairly be called parasitoids. Hosts vary considerably between families. The Embolemidae are poorly understood and little is known about their biology, although they have been reared from Achilidae (planthopper) nymphs, and also found in subterranean nests of ants and mammals. Sclerogibbidae are ectoparasitoids of nymphal Embiidina (webspinners), though once again their biology is poorly known.

The Dryinidae use hosts in the Auchenorrhyncha (leafhoppers, treehoppers, and spittlebugs). Female drynids are wingless and have pincer-like chelae on their front legs that are used to grab onto host insects. When a host is captured, she stings it to temporarily paralyze it. Occasionally the host is eaten (and thus killed) but other times the female will deposit an egg within the leafhopper through the soft membrane between bodily segments. The larvae are usually endoparasitic in the beginning of their lifecycle then gradually becoming ectoparasitic, attached to and feeding on the host from the outside. One species of drynid, Crovettia theliae, is completely endoparasitic and also polyembryonic with a single egg developing into up to fifty genetically identical offspring. Similar to the Dryinidae, Bethylidae are parasitoids of lepidopteran (Bethylinae) and coleopteran larvae and pupae (Epyrinae and Pristocerinae).

Family Chrysididae

The Chrysididae, or cuckoo wasps, are not easily mistaken for other wasps. They are usually a metallic green or blue, occasionally with metallic red on the abdomen or parts of the thorax. They are heavily armored, an adaptation that allows them to enter the nests of other hymenoptera. This family contains both parasitoid and kleptoparasitic species (the lifestyles of the latter give rise to the common name, much like cuckoo birds). Kleptoparasitic species lay eggs in the nests of sphecid wasps (such as Sceliphron spp.), as seen in the photo above, and eat the provisions within the nest of the host insect. Parasitic species are ectoparasitoids of the larvae of various bees, sphecid wasps (Sphecidae), potter wasps (Eumenidae), or more rarely, the larvae of sawflies or the eggs of stick insects.

In many cases the females have highly reduced stingers and venom glands, though there is at least one species with a prominent stinger and venom glands that are large even compared to other Hymenoptera. The Chrysididae have ovipositors, but they are distinct than the ones found in the Parasitica and unique in the Aculeata. Chrysidid ovipositors evolved independently from the aculeate stinger into a telescoping egg-laying apparatus that can withdraw into the abdomen. The ovipositor is used to lay the egg into the nest or pupal case of the host insect.

Subfamily Chrysidinae

Forty four genera in three tribes constitutes the subfamily Chrysidinae. The tribe Chrysidini is by far the largest with nearly half of the species. The Elampini tribe is the nest largest, while the Parnopini is very small with only three genera worldwide. Parnopine wasps prefer arid habitats and use sand wasps (Bembicinae) as hosts. Entry is gained into the nest or puparium of the host by chewing a hole, followed by the deposition of a single egg, then finalizing the process by sealing the hole with debris and saliva, usually of the same material as the structure was originally created.

Tribe Chrysidini

Chrysis angolensis, found dead in a nest of Sceliphron caementarium

Chrysis angolensis is an Old World species that is thought to have been introduced into the United States during or after World War II. They are nest parasitoids of the prepupae of sphecid and crabronid wasps (particularly Sceliphron caementarium). This specimen was found dead in a mud nest of S. caementarium, and is the same individual seen in the previous photo. I had also discovered a number of pupal sacs which most likely belonged to this species. They are a metallic green, but depending on where the light hits it can make it appear blue. This cuckoo wasp probably gained access to the cells with her mandibles to lay her eggs, then sealed the nest again with saliva and debris created when creating the opening.

Tribe Elampini

Omalus sp.? (October)

The tribe Elampini contains around twenty genera of very small chrysidids. They all use other Hymenoptera as hosts, particularly ground nesting wasps in the Crabronidae. Omalus spp. use aphid wasps (Pemphredoninae) as hosts, though other ground nesting wasps are recorded hosts of other elampines such as Hoplisoides spp., Solierella spp., and Tachysphex spp. Old World elampines are also known to use bee hosts in the Halictidae and Megachilidae as well as crabronid wasps.

Omalus sp.? (October)

I watched this tiny wasp scuttling about on my porch stairway, continuously antennating the wood. At first I nearly mistook it for a small dipteran, though a closer look revealed otherwise. This tiny wasp was no longer than around one eighth of an inch (or around three to four millimeters long).

Superfamily Pompiloidea

The superfamily Pompiloidea is probably best known for the spider wasps, Pompilidae, and the so called velvet ants (or "cow killers"), in the family Mutillidae. Most are ectoparasitoids (or occasionally kleptoparasites) of immature Hymenoptera or paralyzed adult spider hosts. Rarely, other insect orders are used as hosts.

Female velvet ants (Mutillidae) are conspicuous at times since they are wingless and meander along the ground searching for hosts or host nests. While the Pompilidae always use spider hosts, the Mutillidae use a variety of hosts from (usually) ground nesting bee and wasp larvae as well as (though rarely) immature stages of flies, slug moths (Limacodidae), beetles, and cockroaches. Female mutillids are the most often encountered, while males are less frequently seen. Unlike the females, the males are winged. Both will visit flowers for nectar, but also feed on honeydew and occasionally females will feed on host insects. Mutillidae differ from other wasps in the Pompiloidea by being very densely hairy and often brightly colored (usually red).

Lesser known and less common families are the Sapygidae and Myrmosidae wasps. Sapygid wasps are parasitoids or kleptoparasites of other aculeate larvae and are particularly fond of twig-nesting Megachilidae bees. Myrmosid wasps are parasitoids of the larval stages of various solitary bees and wasps, and have wingless females. Both of these families are easily distinguished from the Mutillidae since their bodies appear nearly hairless.

Family Pompilidae

The spider wasps are some of the most well known of the solitary wasps. They are also unique among the Pompiloidea in that they always use adult spiders as hosts, either in the spiders own den or in a nest they excavate themselves. Many pompilids seem to use specific genera as hosts. In any case, when a spider is found the wasp will entice it to exit its den and grapple with it until it can sting the spider. Sometimes repeated stings are necessary but paralysis is always permanent. Once dragging the spider back into its den or to a specially dug nest, the pompilid lays a single egg on or near the spider and seals the nest. The pompilid larvae will feed on the living spider until it pupates and emerges the following year. Some sphecid wasps, such as the Sceliphrinae, also use spiders as hosts, but the nests of sceliphrines are aerial while those of most pompilids are subterranean.

Subfamily Pepsinae

With 2000 species in over sixty genera worldwide, the most well known of the Pepsinae are the [relatively] massive tarantula hawks in the genus Pepsis. Tarantula hawks are typically large wasps, some with metallic blue hues and often with orange wings. The largest are up to two inches long, and have the most painful sting of any Aculeate wasp, second only to the bullet ant (Paraponera clavata). Pepsis spp. are more common in arid regions of the American Southwest, but other species ranges extend up to Canada. Most visit flowers, and some are considered important pollinators. Pepsinae appear to specialize in hunting spiders in the Mygalomorphae which includes tarantulas and trapdoor spiders.

Tribe Pepsini

Calopompilus pyrrhomelas (September)

Calopompilus spp. are not quite as large as their Southern cousins, but they do reach about an inch in length which is still large as far as solitary wasps are concerned. C. pyrrhomelas specializes in the folding door spiders in the genus Antrodiaetus. When spiders are paralyzed, they are sealed inside their own den along with an egg. Adult Calopompilus aren't known to frequently visit flowers, but they have been observed visiting rabbitbrush (Ericameria nauseosa) for nectar.

Calopompilus pyrrhomelas (September)

Unfortunately, this individual was found dead in a pool of water. Fortunately, I was able to collect it and I still have it for future study (and because I think it is really really cool). I may not have ever encountered this species this year if it had not drowned. Having found this single specimen suggests that there are more of them.

Subfamily Pompilinae

The subfamily Pompilinae constitutes the largest subfamily in America north of Mexico. The majority of these wasps are completely black, with translucent to dark wings. A few exceptions are Tachypompilus species which are mostly red and Poecilopompilus which are mostly yellow. Most hunt spiders then bring them to a suitable location for excavating a nest, with the exception of the tribe Aporini which hunt subterranean spiders and use the spider's own burrow to rear their young. In either case an egg is glued to the abdomen and the nest sealed. Prey varies widely, from orb weavers (Araneidae) to wolf spiders (Lycosidae), and there have even been reports of harvestmen (order Opiliones) being used. In fact, spiders from at least fifty genera in fifteen families have been recorded as hosts for Pompilinae wasps.

Adults often visit floral or extrafloral nectaries for sustenance, or sometimes seek out honeydew from sap sucking insects. Some species feed on the hemolymph of spider prey, occasionally hunting prey solely for the purpose of feeding on it. This kind of feeding has been demonstrated to improve egg laying and longevity for some wasps.

Tribe Pompilini

Arachnospila sp.? (September)

This spider wasp, and others like it, was seen exploring the outside of an active beehive. It is unclear what its intentions were, though there is a possibility that the scent of honey could have attracted it as many other insects are attracted. Spiders are also very common residents of the undersides of these particular hives, so it is possible this wasp was actively hunting. It disappeared under the lip of the lid shortly after this photo was taken.

Due to the similarity of features between many of the Pompilinae wasps, identification is not simple or easy. Features are probably best examined under a microscope by an expert, which I am not, rather than from inspection of a photo. Nevertheless, much of the information presented here remains the same.

Superfamily Vespoidea

Vespula pensylvanica (left) and Polistes dominula ♂ (right) eating sugar

This superfamily is best known for the social members, particularly paper wasps (i.e. Polistes spp.), yellowjackets (i.e. Vespula spp.), and hornets (Vespa spp.), although there are also many solitary species and obscure rarely encountered species. This superfamily was once considered to contain

While many of the social species are considered to be predatory, there are also parasitoid species which utilize a living host. One mysterious family with no common name, the Rhopalosomatidae, contains a total of four genera worldwide. Wasps in the Rhopalosomatidae, or alternatively rhopalosomatids (the closest we can get to a common name), are ectoparasitoids of cricket nymphs in the subfamily Hapithinae and are thought to be nocturnal.

Family Vespidae

The Vespidae are a large and familiar family with over three hundred species in at least thirty five genera of both solitary, semi-social, and eusocial wasps. The subfamilies Eumeninae, Euparagiinae, and Masarinae are all solitary, Stenogastrinae contains both solitary and social species, Polistinae are mostly semi-social, and Vespinae are completely eusocial. Euparagiinae are very rare and live in arid regions of southwestern United States. Pollen wasps, subfamily Masarinae, provision their nests with pollen rather than insects.

Nest materials vary from paper (plant pulp and saliva) as in the Vespinae and Polistinae to mud (i.e. Eumeninae, Masarinae) to preexisting cavities in wood, the ground, or old nests of other wasps (such as the old mud nests of Sceliphron caementarium). Prey varies as well, and with the exception of the Masarinae they all have carnivorous larvae. Prey are usually larval stages of Lepidoptera but other Hymenoptera are sometimes hunted as well as various life stages of many different types of insects. The Eusocial species seem to display the most diverse dietary habits, and will even take carrion or attack larger adult insects and return it to the nest in small pieces.

Subfamily Eumeninae

I have only encountered one family of solitary vespids, the Eumeninae. Most species, known as mason wasps, nest in preexisting cavities in wood, hollow stems, or in rock crevices, and use mud or sand to partition cells. Some species nest in the ground (all Pterocheilus, Odynerus, and some Euodynerus), and occasionally species build small mud turrets over the nest entrance (i.e. Odynerus dilectus, Euodynerus annulatus). Other species, known as potter wasps, build free standing mud nests (Eumenes and Zeta). Host insects, normally moth larvae but occasionally leaf-feeding beetle larvae, are paralyzed and sealed in the nest cells to be consumed by the wasp larvae. Adults of some species feed directly on the host's hemolymph, and a few eat entire portions of the host's body itself. Kleptoparasites and parasitoids often prey on eumenine nests, chrysidid wasps, mutillid wasps, and sarcophagid flies being the most common of these.

Eumenes verticalis ♀ (October)

Potter wasps are similar in appearance to their eusocial relatives (Vespinae) though they have a unique form, notably the unique shape of the abdomen. I found this female feeding on the nectar of rabbitbrush (Ericameria nauseosa) for at least the duration of my observation, perhaps twenty minutes.

Eumenes sp. nest

Potter wasps themselves are about as recognizable as their nests. Mud nests are attached to plant stems, trees, or sometimes other things like under pallets or other protective structures. The opening is smaller than the wasp itself, and the egg is laid once nest is complete by the female wasp sticking her abdomen through the opening. Normally a single caterpillar is used as a host, paralyzed, and stuffed into the nest and sealed. Unlike sphecid wasps, Eumenes does not need a source of mud nearby and instead can make their own mud by bringing water to mineral soil.

Eumenes sp. prepupae removed from a nest

By autumn, Eumenes larvae have reached the prepupal state (having consumed most of the host) and enter a state of diapause for the winter. The inside of the mud nest is coated with a cellophane-like secretion which serves to strengthen the thin mud walls and may also give it some resistance to water. Most nests are attached in protected locations such as under rocky overhangs or on plant stems.

Ancistrocerus sp. on rabbitbrush (October)

Ancistrocerus, along with Stenodynerus and a number of other genera, are collectively known as mason wasps rather than potter wasps since they use mud or sand to partition cells in tunnels in plant stems, wood, cracks in wood, or in subterranean excavations. Nests are sometimes made in preexisting excavations from other insects, or are even occasionally stolen from bees or other wasps. Some species build small mud turrets at the nest entrance, this is restricted to species which nest in the ground. I have even found old drawn comb of honeybees with mason wasp larvae inside, capped and lined with mud.

Stenodynerus sp. on Levisticum officinale (June)

Stenodynerus is the largest genus north of Mexico with forty eight species. Ancistrocerus has thirty one described species, seventeen in North America. All mason wasps mass provision their nests with caterpillars. They enter winter as prepupae and pupate in the spring.

Subfamily Polistinae

Polistes aurifer

The Polistinae has some familiar members, namely the paper wasps whose nests are typically composed of exposed paper cells in cavities or under overhangs. The common name, paper wasp, is a misleading term since wasps in the Vespinae also have paper nests. Unlike the Vespinae, Polistes and Mischocyttarus have no true worker castes, and all females are fertile (so technically there is no queen, per se). To keep order, there is normally a dominant laying female who will destroy the eggs of "rogue" layers. Brachygastra, however, exhibits complete eusociality with multiple queens in large paper nests and a sterile worker caste.

Similar to other eusocial or semi-social Hymenoptera in the United States, only newly mated females survive over the winter where it is cold. Females find a protected location, such as in a wood pile, to wait out the winter. Upon the onset of spring, females (or foundresses) build small hanging nests under eaves or other protected locations. Paper wasps typically prey on caterpillars but they may take larvae of other insects as well. The reason caterpillars are most often hunted is probably proportionate to their level of exposure and abundance compared to the larvae of other insects. Adults also feed on nectar, and in the genus Brachygastra nectar is brought to the nest and made into honey (thus their common name, honey wasps).

Nests of Polistes and related genera are parasitized by gregarious ichneumonid wasps in the genus Pachysomoides. Moths in the genus Chalcoela (Crambidae) are parasitoids of Polistes and Mischocyttarus nests. Chalcoela caterpillars feed on the wasp larvae, which I find to be ironic.

Mischocyttarus flavitarsis ♀ (October)

Mischocyttarus is very similar to Polistes except for the elongates and distinctively curved petiole. It is known as the Western paper wasp due to its westerly distribution. I spotted this female while working some beehives, honey had gotten onto the lid of an adjacent hive and she was feeding on the honey. Paper wasps aren't interested in hunting honeybees, but they have been known to rob hives of their resources. Another interesting adaptation I have observed is the ability for some paper wasps, particularly those with very long legs, to land directly on still water. Surface tension allows them to alight, drink, and take off without drowning.

Mischocyttarus flavitarsis ♂ (November)

A male Mischocyttarus on my finger. Males of many vespid wasps can be identified by their curled antennae. The green eyes are also distinctive from females, and the faces of some species are unmarked yellow. Males have no stinger, so there is no risk when holding one.

Polistes aurifer ♀ (November)

A female Polistes aurifer, known as the golden paper wasp, on my finger. I found this female hiding on some empty beekeeping equipment, her winter shelter. She was cold, moving slowly, and she let me photograph her for a while. Although she has a stinger and was fully capable of stinging me, I gave her little incentive to do so. Note that she is holding me, not the other way around.

Polistes aurifer ♀ (July)

While working in one of the apiaries this year, I had discovered a nest underneath some empty bee equipment. There were two or three workers who fled while the dominant female was unwilling to leave. While I had my beekeeping veil on, I decided to get some photos. I question whether or not I even needed a veil since there were no signs of aggression. She appeared to be missing part of her right forelimb, possibly in a battle for dominance with another female. Body size seems to be a determining factor with nest dominance, a result of good nutrition. Well fed larvae grow up to become larger adults, and larger adults can defeat smaller opponents.

Polistes aurifer eggs (July)

Nests of all paper wasps are made from plant and wood fibers held together with saliva, but they appear to use whatever is available including construction paper or even money. Unlike honeybees, many of the social vespid wasps (at least American species) build horizontal comb with the cells facing down. Paper wasps typically build a single rounded comb.

Polistes aurifer ♂ on rabbitbrush (October)

Males are produced towards the end of summer or the beginning of fall. They will mate with new females once and die. Like the Mischocyttarus male, the face is completely yellow with antennae that curl slightly at the tip.

Comparison between Polistes aurifer (L) and P. dominula (R), not to scale.

A common European invader in much of the United States, the European paper wasp (Polistes dominula, pictured above on the right) is slightly smaller than P. aurifer (photos not to scale). This species is slightly more aggressive, and often shares habitat with P. aurifer (which in my case is under the eves of my roof). They can be told apart by their patterns, higher contrasting patterns of P. dominula while the abdomen of P. aurifer is mostly a dull yellow. I would also like to add the observation that the antennae differ in color, P. aurifer antennae fade to black at the ends while P. dominula antennae are black at the base. The yellow markings on the thorax of P. dominula are also diagnostic.

Polistes dominula ♀, missing her right forelimb. (October)

The European paper wasp is native to Europe and parts of Asia and Africa and was first recorded in North America in 1981. It has been a highly successful colonizer and it expanded its range in the eastern and midwestern USA in the twenty years following its introduction. As an invasive species, Polistes dominula directly competes with native paper wasps, P. aurifer and Mischocyttarus flavitarsis. P. dominula establishes colonies earlier than the native species, and can thus get a head start on finding the most suitable nesting sites. Smaller body sizes than the native species, coupled with their preference for small holes and voids allows them to nest in a greater variety of nest sites than the larger bodied native species. Old or empty nests left over from the previous year may be reused by P. dominula, while native species do not. While the native species are primarily caterpillar hunters, European paper wasps may hunt a variety of insect types from many orders, including the larvae of other wasps or even other paper wasp nests.

Polistes dominula ♂ on my hand, males are incapable of stinging yet he did poop on me. (October)

Male P. dominula wasps have the conspicuously curled antennae and mostly yellow face. Like all the other eusocial or semi-social vespid wasps, they are produced in late summer or early fall to mate with new females who will overwinter and form new colonies in the spring.

Polistes dominula ♀ (October)

Some, but not all, Polistes wasps use a form of facial recognition to communicate with nestmates and understand hierarchy. Species with less descript facial patterns do not communicate this way, and instead have comparatively larger olfactory sensors. P. dominula and P. fuscatus are two that have demonstrated facial recognition of nestmates. There is also some speculation that they can recognize the faces of people, though I could not find anything to substantiate this.

Subfamily Vespinae

The subfamily Vespinae contains the most infamous of the social wasps: yellowjackets and hornets. They are markedly more aggressive and protective than other wasps, and their preference for nesting in trees and cavities (subterranean or otherwise) often places them near people. Nests of many yellowjackets and hornets can become very large, excessively large where winters are mild and colonies can perennialize. They are truly eusocial, having distinct castes complete with reproductive (queen, drone) and worker (sterile female) castes.

Much of the bad press comes from this group of wasps. They are also the most likely to turn up at a picnic or barbeque. Nest sites often overlap with human activity. In these ways they often interact with people. If a nest is approached or accidentally disturbed (such as mowing the grass and being unaware that a yellowjacket nest is underfoot), the unsuspecting human is met with an aggressive or at the very least intimidating response. This is bad for obvious reasons, but this is in fact how these creatures have been able to survive for millions of years. A colony is full of delicious treats, particularly if you are a bear (or one of a number of other mammals), and the highly aggressive response is their best means of defense only second to their typically hidden or inaccessible nest sites.

Vespula pensylvanica queen feeding on nectar from plum blossoms. (March)

Yellowjackets are one of the most commonly encountered social wasps in the United States. In the South they can form massive colonies, both above- and below ground, since they can perennialize through the mild winters. In temperate America, they form annual (occasional perennial) subterranean colonies. Newly mated queens are the only ones to survive cold winters, similar to the paper wasps (Polistinae). Upon emergence in the spring, they seek nest sites and feed on nectar from flowers. Surely they contribute to pollination, some species are quite hairy and fully capable of inadvertently spreading pollen.

Similar to honeybees, yellowjackets (Vespula spp.) utilize pheromones as one means of communication. Queen yellowjackets produce pheromones that induce sterility in the female worker caste, preserving her ability to preserve the queens genetic lineage. Another result of yellowjacket queen pheromone enables workers to recognize her eggs versus the eggs laid by workers. Laying worker eggs are destroyed or eaten.

Vespula pensylvanica ♂ (October)

Male yellowjackets have the distinctive and highly elongated antennae. They are likely used to search for virgin queens to mate with. Yellowjacket drones are produced late in the year, early autumn or late summer. Similar to honeybee drones, they mate once and die, their genitals torn from their body resulting in death.

Vespula pensylvanica worker (August)

Yellowjacket workers are the workhorses of the colony. Yellowjackets are omnivorous, as adults may feed on flowers, but carnivorous as larvae. Yellowjackets feed on a variety of other creatures, from insects of many orders to carrion to hamburgers. They will attack insects much larger than themselves and bring it back in small chunks to feed to their young. They will also rob resources from honeybee hives, much to the detriment of the hives and the beekeeper. In my experience, a strong hive should adequately fend off even the largest of yellowjacket populations while small colonies may be in trouble.

Vespula pensylvanica devouring a honeybee (August)

Yellowjackets are a major pest in an apiary. They will attack honeybees in the air or go right into the hive to attack them. They may also play a role as scavengers, bringing home bees which have died of other causes. In my experience, keeping one's hives at their peak health and utilizing reduced entrances can go a long way in mitigating the issue of yellowjackets.

Dolichovespula maculata ♀

Bald-faced hornets, technically a species of aerial yellowjacket, form large watermelon-sized nests which hang from the branches of trees. Their nests are composed of multiple levels of horizontal paper combs with an outer shell composed of many sheets of paper. The stereotypical image of a beehive hanging from a tree branch is more typical of the nest of Dolichovespula maculata.

Dolichovespula maculata ♀

These are large wasps, and create colonies with seven hundred or more individuals. They have diets similar to yellowjackets (Vespula spp.) and also commonly hunt honeybees where both are common. Honeybees are smaller, but have various defensive strategies to fend off or fight larger foes. When stinging doesn't work, they will form a tight ball of many individual bees around an attacker and vibrate their bodies to heat it up. So much heat is generated that they kill the attacker. This was probably the case with this individual. It is dead, and I found it fully intact at the bottom of a beehive earlier this year.

Dolichovespula maculata ♂ with Apis mellifera forager (October)

Similar to all other social vespid wasps, males are produced in the fall. Males were a common sight in October. I was working some hives when this male (note the curled antennae) landed on my glove. A worker honeybee landed next to him to investigate, then departed after losing interest.

Parasitica

Parasitoid wasp (Lissonota sp.) with a long ovipositor used for penetrating hosts and laying eggs.

The Parasitica is composed largely of obscure, rarely encountered solitary wasps that use
(usually) living hosts to raise their young. The primary distinction between the Aculeata and the Parasitica is the evolution of a stinger from an ovipositor. The Aculeata evolved from the Parasitica, it is believed, and lost the use of their ovipositor for laying eggs in exchange for a stinger capable of stinging and injecting venom. For the Aculeata, eggs are dropped from an opening at the tip of the abdomen under the stinger. The Parasitica, however, deliver eggs with an ovipositor. Some have relatively short ovipositors, while others have very long ovipositors capable of penetrating solid wood when seeking wood-boring hosts or penetrating soil when seeking soil dwelling hosts. Venom is usually injected along with the eggs, and despite lacking true stingers some parasitoids are capable of using their ovipositor to sting people, though this probably requires provocation as I have discovered these creatures are very quick to flee.

Hosts vary considerably, in nearly every insect order and even other arthropods such as spiders and mites. Eggs, larvae, prepupae, pupae, and adult hosts are used depending on the species of parasitoid wasp. Some wasps use plants as hosts, such as gall wasps (i.e. some Chalcidoidea and all Cynipoidea). Eggs can be laid near, on, or within a host, and larval development can take place on or within a host at different stages depending on the species. Some members of the Parasitica are known as hyperparasitoids, those which attack the parasitoid larvae within another host (thus unintentionally saving the original host in some cases).

Polyembryonic parasitoids are those that lay one egg inside a host which develops into many larvae, and depending on the species many wasp larvae can emerge from a single egg laid, or even a few thousand in the genus Copidosoma! The eggs of polyembryonic species occasionally divide within a host or there may be multiple embryos within a single egg. Many other fascinating scenarios take place, far beyond my scope of knowledge. To read more about the Parasitica I suggest these books: Bees, Wasps, and Ants by Eric Grissel (Timber Press 2010), Parasitic Wasps by Donald L. J. Quicke (Springer 1997), and Solitary Wasps by Kevin M. O'Neill (Comstock Publishing 2000).

Superfamily Chalcidoidea

The Chalcidoidea is a large and varied group of small wasps with an incredibly diverse array of lifestyles. There are well over 500,000 species described worldwide, and over 2,000 in the United States alone. Most are parasitoids of insects and arachnids, yet some are plant tissue feeders who grow up within stems, leaves, seeds, or flowers, or even make galls. Fig wasps, family Agaonidae, develop and mate entirely within figs. Some are parasitoids of pest insects (within the orders Lepidoptera, Diptera, Coleoptera, and Hemiptera) and are used for pest control purposes. At least one species, Tetramesa romana (Eurytomidae), has been released into the US to control an invasive grass, Arundo donax (Poaceae).

The smallest wasps, and smallest insects, are in the families Mymaridae and Trichogrammatidae, both contain egg parasitoids of very small host insects: Thysanoptera (thrips) and Psocoptera (barklice). The most peculiar species, however, are those which are hyperparasitic and use other parasitoids as hosts. This can be in the form of secondary or tertiary parasitoids, or to the extreme as quaternary parasitoids. This equates to a parasitoid parasitizing a parasitoid parasitizing a parasitoid of some sort of unfortunate host. It is fathomable that such an interaction may even save the original host, but only if the secondary parasitoid parasitizes the primary parasitoid before pupation, the point at which the original host would be severely weakened by the primary parasitoid. Crystal clear, right? Other species may be polyembryonic.

In all, chalcid wasps attack insects in around 340 families representing fifteen orders, as well as some arachnids (i.e. pseudoscorpions, ticks, and mites). Mostly immature stages of hosts are parasitized, but adult hosts are sometimes utilized as well as eggs and egg sacs (including spider egg sacs, in some cases).

Chalcid wasps hovering around Quercus kelloggii (April)

I took this photo to capture the light shining through the new growth of the California black oak, and only later did I discover the wasps. Expert identification suggested Chalcidoidea, though there is no way to be sure so allow me to speculate. Oak trees are host to many insects, including many species of moths which may play the role as hosts to countless species of parasitoid wasps. Gall wasps, Cynipoidea, are also found to associate with oaks in abundance. All of these insects are possible hosts for chalcid wasps, depending on the species.

Superfamily Cynipoidea

Family Cynipidae

The family Cynipidae is an oddity in the world of wasps (though if you have read this far, it has probably come to light that wasps, particularly the Parasitica, are an oddity as a whole). Cynipid wasps are mostly gall forming, phytophagous (herbivorous) wasps which induce galls in which to feed on and live within during their development. Inquilines don't make galls themselves, but feed on the gall of another wasp, occasionally killing the host larvae.

Gall forming wasps initiate the formation of a gall when certain chemicals excreted by the wasps make contact with the plant tissue, though the exact mechanism is unknown in most cases. The type of plant or tree as well as the location of the gall is very specific to the species of wasp. Most gall wasps only form galls on a select few or single species on plant or tree. Usually oaks (Quercus spp.) and other trees of the family Fagaceae, or roses (Rosa spp.) are most often used as host plants for gall formation. There are some herbaceous gall formers who favor plants in the families Asteraceae, Lamiaceae, and Papaveraceae, though their galls are less conspicuous.

The gall serves two primary purposes for the wasps: as a source of food for developing larvae and to protect the immature wasp from its natural enemies. The inside of the gall is full of nutritious plant tissue, and a source of protein for the wasp larvae. The outside of the gall is often hardened to form a shell-like exterior. Sometimes the outer layers are toxic, a protective measure that keeps predators from eating the gall.

There are at least fourteen hundred described species of cynipid wasps in the world, 750 in around fifty genera in North America. Many have interesting sex lives. Some species are believed to be completely asexual with completely female populations. Others have strange reproductive cycles. Some bivoltine species (two generations per year) have generations which alternate between sexual and asexual (parthenogenetic), such that the sexual generation which has both male and female members will yield only females. The asexual (entirely female) generation will not mate, yet will yield both male and female offspring. Quite alien when compared to our human lives!

Tribe Cynipini

Andricus quercuscalifornicus gall

By far the largest and most conspicuous galls in the West, the large oak apple galls formed by the wasp Andricus quercuscalifornicus are familiar to many who live among oaks. Female wasps lay eggs on stems/twigs of various oak species. After the eggs hatch, the gall develops around the tiny larvae. Galls become very large, and occasionally multiple eggs are laid so the gall may support many larvae. Only parthenogenetic (asexual) females are known from this species.

Andricus quercuscalifornicus gall interior

Many other insects utilize the large galls or parasitize the larvae of the common Andricus quercuscalifornicus wasps. Inquilines include the moth Cydia latiferreana (Tortricidae) whose larvae feed on the inside of the gall. The moth larvae is then used as a host to the parasitoid wasp Bassus nucicola (Braconidae). Beetles (Ozognathus cornutus, Anobiidae) and termites (Rhinotermitidae) have also been found feeding on the gall material. Chalcid wasps in the families Eulophidae, Eurytomidae, and Torymidae have been documented as parasitoids of A. quercuscalifornicus, though surely there are many more inquilines and parasitoids than we know of.

Cynips mirabilis gall on Quercus garryana leaf (October)

A smaller yet more ornate gall is that of the gall wasp Cynips mirabilis. I found several small galls, perhaps twelve to thirteen millimeters in diameter, on the undersides of the leaves of Quercus garryana. It is probably safe to infer that eggs are laid sometime in the spring as leaves emerge, and development must occur within the year before the leaves drop in the fall.

Cynips mirabilis gall opened

Within the outer shell, a small hardened mass (which houses and feeds the larva) is suspended by delicate filaments. This gall is occasionally host to inquilines, since the larvae of moths (as yet unidentified) have been found within these galls.

Tribe Diplolepidini

Diplolepis rosae galls on a wild rose plant

As the specific epithet implies, Diplolepis rosae forms galls on a select few species of roses. D. rosae is a European native, accidentally introduced into the United States. Wasps lay eggs on the young leaf buds in spring as new leaves emerge. Multiple larvae feed on a single gall (personal observation), and they remain inside the gall throughout the winter before pupating and emerging as adults in spring. They are possibly parthenogenetic in some populations, as very few males have ever been documented. Ichneumonidae and Chalcidoidea wasps have been known to parasitize the larvae of D. rosae.

Superfamily Diaprioidea

Family Diapriidae

Subfamily Diapriinae

Aneurhynchus sp. ♂

Aneurhynchus spp. are known to parasitize flies which feed on decaying plant matter, such as Fannia spp. (Fanniidae), Platypeza spp. (Platypezidae) and flies in the Sciaridae. These flies feed on either decaying plant material or fungus, mostly in wooded areas.

Aneurhynchus sp. ♂

I encountered this tiny wasp (~3mm) at my porch light at night. I imagine these are very common, yet are very often disregarded as small flies.

Superfamily Evanioidea

The Evanioidea contains three families: The Aulacidae, Evaniidae, and Gasteruptiidae. The Aulacidae and Gasteruptiidae are described below, but I have not encountered the Evaniidae, sometimes known as ensign wasps. Ensign or hatchet wasps (Evaniidae) are single egg predators within the egg cases, or oothecae, of cockroaches. Evaniids are peculiar in appearance for their much reduced abdomens, in contrast to the characteristic abdomens of the other two related families. There are at least 1,100 known species, probably many more, worldwide.

Family Aulacidae

The Aulacidae are a group of parasitoids often mistaken for ichneumon wasps which often share similar habitats and host preferences. Aulacids are often found in wooded areas. Aulacid wasps seek out wood boring larvae of beetles (several families, but mostly longhorns, Cerambycidae) and wood wasps (Xiphydriidae, Symphyta). Female Aulacids antennate the wood, seeking the host, and once a host is detected (perhaps they can sense the subtle movements of wood boring larvae?), they insert their ovipositor through the bark and into the host, where an egg is deposited.

Pristaulacus sp. ♂ (August)

I discovered a male Aulacidae, evident by the lack of the long ovipositor, struggling in a pool of shallow water. I attempted to rescue him in my hand where he proceeded to groom himself for about ten minutes before unexpectedly perishing.

Pristaulacus sp. ♂ (August)

One unique feature of this group of wasps is the high attachment point of the abdomen, something shared with the Gasteruptiidae which the Aulacidae is closely related. Also of personal fascination are the ridged on the top of the thorax. Aulacids are known to occasionally visit flowers for nectar, though I have not observed this directly.

Family Gasteruptiidae

The family Gasteruptiidae have unique and easily identifiable abdomens, though little is known by the lay observer of their biology. They are sometimes known as carrot wasps, probably a hint of the forage preferences of adult wasps seeking nectar, though they will visit other plants with small flowers that have easily accessible floral resources.

Gasteruption sp. ♂ on Allium cepa (June)

Larvae of gasteruptiid wasps are predators or predator-inquilines of other Hymenoptera that nest in pithy stems or twigs of woody plants. Predator-inquilines eat the larval food in the nest rather than the larvae itself, while predatory gasteruptiids feed on the larvae itself, such is the case for Gasteruption spp. Either way, the host larvae dies. Gasteruptiids go through winter as prepupae, then pupate in the spring.

Superfamily Ichneumonoidea

Over 6,000 species described north of Mexico, and at over 100,000 worldwide make up the superfamily Ichneumonoidea. The family has two major families: the Braconidae and Ichneumonidae. Both utilize a diversity of hosts across many insect orders and even some arachnids in various life stages from egg to adult.

The most peculiar feature among some of the Ichneumonoidea (notably the subfamilies Campopleginae and Banchinae in the Ichneumonidae, and subfamilies Microgastrinae, Miracinae, Cheloninae, Adeliinae, Cardiochilinae, Khoikhoiinae, and Mendesellinae in the Braconidae) is the presence of polydnaviruses in their genomes. Polydnaviruses are released into the host when an ichneumonoid is ovipositing, serving to suppress the immune system of the host and thus protecting the egg. The polydnavirus then infects the developing wasp larvae. Specific polydnaviruses have coevolved with the wasps they infect, and the wasps have evolved to be dependant on the virus to protect the developing wasp larvae.

Family Braconidae

Subfamily Alysiinae

About 2,300 described species make up the subfamily Alysiinae. They are small wasps, generally less than five millimeters but usually half that. While adults may feed on nectar and possibly pollen, they use various families of flies as hosts for their immature stages, mostly flies in the Agromyzidae, Anthomyiidae, Calliphoridae, Chloropidae, Ephydridae, Phoridae, and others. Female alysiine wasps lay their eggs in the eggs or larvae (maggots) of host flies, but don't complete their development until the host enters the next phase of development (either as larvae or pupae, respectively). In the case of larval-pupal parasitoids (those that parasitize the fly in the larval stage), they use specially adapted inverted mandibles to cut a hole through the hardened fly puparium instead of the mature fly.

Alysiinae (October)

This wasp, probably less than three millimeters in length, was attracted to my porch light in the evening. Many small dipterans were also attracted to the lights, some may even been hosts for this small wasp but I never attempted to identify them.

Subfamily Aphidiinae

Aphidiine wasps are not as numerous of the aforementioned wasps, yet are probably common where they are found. Around 120 species in sixteen genera inhabit the United States, and only 400 species in fifty genera are described around the world. They are endoparasitoids of aphids, which are only partially paralyzed before an egg is deposited. The host continues to develop while the tiny wasp larvae eats it from the inside, but dies as the wasp approaches the pupal stage. Aphidiines pupate within the empty shell of the host aphid after eating everything inside.

Aphidiine wasp (right) with a crane fly (Tipulidae, left)

Aphidiines are small wasps, not much larger than three millimeters in total length. The photograph I took is misleading, the crane fly in the background is much larger than the tiny wasp. Like some of the other small parasitoid wasps I've encountered, it was drawn to the light of my porch light.

Family Ichneumonidae

The Ichneumonidae, or ichneumon wasps, are endo- or ectoparasitoids of a wide variety of insects and arachnids (including immature or adult spiders, and egg sacs of spiders and pseudoscorpions). Their hosts are always insects which exhibit complete metamorphosis (where an insect goes through four distinct stages: egg, larvae, pupae, and adult) rather than incomplete or gradual metamorphosis. They often but not always have elongated ovipositors, and are partially distinguished from the braconids by a vein structure on their forewings which resembles a horse head, as well as a number of other anatomical structures. Like most Parasitica, they are incredibly difficult to identify and any attempt at identification should really be left to an expert.

Subfamily Banchinae

Tribe Atrophini

The tribe Atrophini uses lepidopteran (occasionally coleopteran) hosts, usually moths in the family Tortricidae, though they also parasitize moths in the families Pyralidae, Cossidae, Sesiidae, Gelechiidae, and others in both concealed and exposed locations. For these reasons, the genera Australoglypta spp., Cephaloglypta spp., Glypta spp. and possibly others have been used in biological control efforts.

Cryptopimpla amblipennis (October)

Cryptopimpla amblipennis parasitizes larvae or prepupae of both beetles and moths (families Erebidae, Lasiocampidae, and Saturniidae) with short ovipositors. Development of the wasp is completed when the host enters the pupal stage and has built a cocoon.

Cryptopimpla amblipennis (October)

Like a few the other wasps here, this wasp was drawn to the light of my porch light. It flew around then landed on the glass of the lamp and walked around. This was repeated for an hour, at least.

Lissonota sp. ♀ (October)

Lissonota spp. are endoparasitoids of Lepidoptera in concealed locations, probably in wood. The host is only partially paralyzed during oviposition, then continues to develop. The wasp matures after the moth has built its cocoon.

Lissonota sp. ♀ (October)

I discovered this small wasp (around one centimeter long) on the side of a beehive on a cool day. Perhaps it was attracted to the warmth of the hive?

Subfamily Ophioninae

Despite only having thirty-seven species in nine genera in the US, the Ophioninae are some of the most common ichneumon wasps encountered. Many are nocturnal, and drawn to lights, while others are diurnal or crepuscular. Most are endoparasitoids of lepidopteran larvae, though at least a few parasitize beetle larvae. Most prey on host caterpillars that feed on vegetation such as members of the families Noctuidae, Lasiocampidae, Lymantriidae, Saturniidae, Geometridae, Arctiidae, and Sphingidae. The ovipositor is very sharp (although they've been known to sting humans) and paralysis of the host is temporary.

Ophion sp. (March)

Ophion is a very common genus in both the United States and in the United Kingdom, where they are often drawn to lights at night. Most species are nocturnal, when they seek caterpillar hosts. Usually one egg is laid in each host. Development of the host continues until it reaches the pupal stage when it typically dies while the wasp remains inside to pupate itself.

Subfamily Ichneumoninae

The subfamily Ichneumoninae is made up of both ecto- and endoparasitic wasps which parasitize caterpillars of both lepidopterans and symphytans (sawflies). They have short ovipositors, so must find hosts that are in exposed locations such as external plant feeding caterpillars. Ichneumonine wasps are idiobionts, meaning their hosts cease or slow development after oviposition. Hosts are parasitized typically in the larval or pupal stage, but wasps always emerge from pupae. Adults occasionally eat large portions of the host bodies, sometimes killing them in the process. Adults also feed on honeydew from aphids or other sap sucking insects, and are said to eat the foliage of certain plants themselves.

Tribe Ichneumonini

The tribe Ichneumonini are represented by over 450 species north of Mexico, one third of which are in the genus Ichneumon.

Spilichneumon sp. (March)

Spilichneumon spp. are internal parasitoids of noctunid moth larvae. The caterpillar reaches the pupal stage, then perishes as the wasp continues to pupate and later emerges from the chrysalis. Adult wasps overwinter as adults, then seek prey in leaf litter the following spring. I discovered this wasp on sticky whiteleaf manzanita (Arctostaphylos viscida) where it may have been feeding on honeydew or perhaps the sticky secretion of the plant itself.

Superfamily Formicoidea

Family Formicidae

The monotypic family Formicidae, the ants, are represented by 14,000 species worldwide. They evolved from wasps between 110 and 130 million years ago. Most are eusocial with fully defined castes, or are workerless parasites which require the eusocial medium of a host ant's colony to reproduce and survive.

Ants are differentiated from wasps by the constriction of their waists, which typically have two or three points of constriction rather than the single point of constriction of most wasps. Ants have elbowed antennae, while very few wasps have elbowed antennae, but the wasps that do have elbowed antennae live solitary lives while all ants are social. Ants are typically wingless, except during mating cycles when winged male and female (queen) ants leave the nest to mate with ants from different colonies and form new colonies. Males die after mating, and females lose their wings after they have finished mating. Females may life from one to over twenty years, depending on the species.

Ant diets vary widely. Some feed on fungus they tend to in the nest, others feed on honeydew, plant nectaries (floral or extrafloral), or other insects. Few are considered to be pollinators, although there are some orchids specifically pollinated by ants (i.e. Chamorchis spp., Microtis spp.). The chemicals secreted by ants are damaging to the pollen of some plants, and although ants may be seen visiting a wide variety of flowers for nectar, the pollen that adheres to their bodies may be rendered sterile. Admittedly, it is an understudied topic and there are probably more ant pollinated plants than we are currently aware of.

Subfamily Formicinae

Named for the ant genus Formica (Latin for ant), the Formicinae are occasionally large wasp-like ants complete with a vestigial stinger and venom gland for production and retention of formic acid. Most feed on nectar and honeydew, but some species are also scavengers or predators.

Tribe Camponotini

Camponotus sp. (June)

The carpenter ants (Camponotus spp.) are large and conspicuous ants associated with rotting wood. Despite their name, they don't eat or chew sound wood but rather infest wood that is already compromised from dry rot (fungal) or other causes of decay. Nests can be in wood (fallen logs, or in this case, house siding) or in the ground, rarely in living wood. They are generalists and omnivorous, eating a wide variety of foods including honeydew, sap, living or dead insects, or a variety of other things. In the evenings they are often found to be grouped near the entrance of their nest, and will all eventually move inside.

Subfamily Myrmicinae

The Myrmicinae are highly variable in size, with the smallest being about one millimeter and the largest up to one centimeter. Many species are generalist predators, although many others are seed harvesters. Others still are fungal cultivators, collecting plant material to grow their subterranean fungus crop.

Tribe Attini

Pheidole sp. (August)

Big headed ants, Pheidole spp., are so named for the large heads of some of the workers (the majors, the minors are smaller). Many species of ants exhibit polymorphism, where two or more forms of a single caste have different proportions and with them slightly different tasks. Many species of Pheidole are generalist scavengers which feed on a wide range of prey items, while others are seed harvesters. They were mostly active at dusk when the temperatures began to drop. Harvester ants typically forage in hot dry periods, but they obtain water from metabolizing the fats within the seeds (or other prey items) they gather.

Pheidole sp. (August)

The orange mass is a piece of protein supplement made for honeybees. I was curious if they would have an interest in it as food. The pollen patty was of minimal interest to the ants, who were probably more interested in seeds than the soy and sugars that made up the supplement.

Tribe Stenammini

Aphaenogaster sp. (August)

Aphaenogaster contains seed harvesting, scavenging, and aphid farming species. These are agile, and somewhat large ants (though not as large as Camponotus) with long legs and slender bodies. This helps them steal prey from other predators, being the opportunists that they are. On occasion some species tend to aphids feeding on roots of plants within their nests, the ants themselves feed on the honeydew excreted from the aphids.

Most species of Aphaenogaster nest in the ground among stones or logs, and their nest entrances can sometimes be identified by a ring of pebbles and other debris. They are active at dusk when it is very warm, since they are active primarily in summer in my area. Nests move frequently, perhaps to find new host roots for their aphid associates. I attempted offering them a piece of pollen substitute (made for honeybees) and unlike the Pheidole ants they were apparently quite pleased with it as much of it was eaten.

Aphaenogaster sp. (August)

Last year I documented nine species of wasps from six families. This year I documented at least twenty nine species (including the ants) in at least twelve families. The more I learn about the amazing variety of wasps in the world, or even just around me, the more enthralled I become. Wasps are the progenitors of modern bees and ants, and their contribution to our world and our lives is untold. I am optimistic that by showing my photos and compiling what I have learned and observed about the wasps within my periphery I can show others that wasps are not to be feared or eliminated but celebrated and cherished. Wasps are pollinators, predators, and food for vertebrates (including humans, in some cases). Learn about wasps, make some observations of your own, and report your findings. What could possibly go wrong?

References

Akre, Roger D., and Hal C. Reed. "Evidence For A Queen Pheromone In Vespula (Hymenoptera: Vespidae)." The Canadian Entomologist 115.04 (1983): 371-77.https://doi.org/10.4039/Ent115371-4

Bohart, R. M., and E. E. Grissell. "California Wasps of the Subfamily Philanthinae." Bulletin of the California Insect Survey 19 (1975)http://essig.berkeley.edu/documents/cis/cis19.pdf

Bohart, R.M. & Menke, A. S. 1976. Sphecid Wasps of the World: a Generic Revision. University of California Press, Berkeley, California.

Garvey, Kathy Keatly. "What a UC Davis Research Team Learned About Oak Apple Galls." Department of Entomology and Nematology. UC Davis, 29 Mar. 2011. Web.http://entomology.ucdavis.edu/News/What_a_UC_Davis_Research_Team_Learned_About_Oak_Apple_Galls/

Grissell, Eric. Bees, Wasps, and Ants: The Indispensable Role of Hymenoptera in Gardens. Portland, Or.: Timber, 2010. Print.

Gronenberg, Wulfia, Lesley E. Ash, and Elizabeth A. Tibbetts. "Correlation between Facial Pattern Recognition and Brain Composition in Paper Wasps." Brain, Behavior and Evolution 71.1 (2007): 1-14.https://www.researchgate.net/profile/Elizabeth_Tibbetts/publication/5963821_Correlation_between_Facial_Pattern_Recognition_and_Brain_Composition_in_Paper_Wasps/links/55d7868e08ae9d65948d9416.pdf

Hammon, Bob University. "The European Paper Wasp." Colorado State University Extension, 26 Apr. 2014.http://www.wci.colostate.edu/shtml/EuropeanPaperWasp.shtml

Johnson, Brian R., Marek L. Borowiec, Joanna C. Chiu, Ernest K. Lee, Joel Atallah, Philip S. WardPhylogenomics Resolves Evolutionary Relationships among Ants, Bees, and WaspsCurrent Biology, Volume 23, Issue 24, 16 December 2013, Page 2565http://www.sciencedirect.com/science/article/pii/S0960982213010567

Joseph, Maxwell B., Melanie Gentles, and Ian S. Pearse. "The Parasitoid Community of Andricus Quercuscalifornicus and Its Association with Gall Size, Phenology, and Location." Biodiversity and Conservation 20.1 (2010): 203-16.

LaSalle, J., and I. D. Gauld. "Parasitic Hymenoptera and the biodiversity crisis." Redia 74.3, Appendix (1991): 315-334.
Losey, John E., and Mace Vaughan. "The Economic Value of Ecological Services Provided by Insects." BioScience 56.4 (2006): 311.http://bioscience.oxfordjournals.org/content/56/4/311.short

Noyes, J. S. "About Chalcidoids." Universal Chalcidoidea Database. Natural History Museum, June 2003.http://www.nhm.ac.uk/our-science/data/chalcidoids/introduction.html

O'Neill, Kevin M. Solitary Wasps: Behavior and Natural History. Ithaca: Comstock Pub. Associates, 2001. Print.

Oi, Cintia A., Annette Van Oystaeyen, Ricardo Caliari Oliveira, Jocelyn G. Millar, Kevin J. Verstrepen, Jelle S. Van Zweden, and Tom Wenseleers. "Dual Effect of Wasp Queen Pheromone in Regulating Insect Sociality." Current Biology 25.12 (2015): 1638-640. Web.http://www.cell.com/current-biology/fulltext/S0960-9822(15)00491-1

Owen, T. L. "Croton Setigerus." Croton Setigerus. Amateur Anthecologist, 24 Aug. 2016.http://www.amateuranthecologist.com/2016/08/croton-setigerus.html

Owen, T. L. "Polistes Aurifer." The Amateur Anthecologist, 8 Nov. 2016.http://www.amateuranthecologist.com/2016/11/polistes-aurifer.html

Owen, T. L. "Sceliphron Caementarium.The Amateur Anthecologist, 13 Aug. 2016. Web.http://www.amateuranthecologist.com/2016/08/sceliphron-caementarium.html

Owen, T. L. "Wasps of 2015." The Amateur Anthecologist. 13 Dec. 2015.http://www.amateuranthecologist.com/2015/12/a-year-of-pollinators-wasps.html

Poinar Jr., G. O. & B. N. Danforth (2006). "A fossil bee from Early Cretaceous Burmese amber". Science. 314 (5799): 614.
Polilov. 2011. The smallest insects evolve anucleate neurons. Arthropod Structure and Development http://dx.doi.org/10.1016/j.asd.2011.09.001

Radisics, Milan. "Hidden Life of the Beewolf." Maptia. N.p., May 2016.https://maptia.com/milanradisics/stories/hidden-life-of-the-beewolf

Regier, Jerome C., Jeffrey W. Shultz, Andreas Zwick, April Hussey, Bernard Ball, Regina Wetzer, Joel W. Martin, and Clifford W. Cunningham. "Arthropod Relationships Revealed by Phylogenomic Analysis of Nuclear Protein-coding Sequences." Nature 463.7284 (2010): 1079-083.

Ronquist, Fredrik, José-Luis Nieves-Aldrey, Matthew L. Buffington, Zhiwei Liu, Johan Liljeblad, and Johan A. A. Nylander. "Phylogeny, Evolution and Classification of Gall Wasps: The Plot Thickens." Plos One 10.5 (2015).http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0123301

Sheehan, M. J., and E. A. Tibbetts. "Specialized Face Learning Is Associated with Individual Recognition in Paper Wasps." Science 334.6060 (2011): 1272-275.http://science.sciencemag.org/content/334/6060/1272

Steinberg, Avi. "The Connoisseur of Pain." The New York Times. N.p., 18 Aug. 2016.http://www.nytimes.com/2016/08/21/magazine/the-connoisseur-of-pain.html

Wasbauer, M. S., and L. S. Kimsey. "California Spider Wasps of the Subfamily Pompilinae." Bulletin of the California Insect Survey 26 (1985)http://essig.berkeley.edu/documents/cis/cis26.pdf

Yong, Ed. "How Tiny Wasps Cope with Being Smaller than Amoebas | Not ..." Discover. Kalmbach Publishing Co., 30 Nov. 2011.http://blogs.discovermagazine.com/notrocketscience/2011/11/30/how-fairy-wasps-cope-with-being-smaller-than-amoebas/

"Family Group Names and Classification [of Sphecidae Sensu Lato]." Hymenoptera. Wojciech J. Pulawski comp. California Academy of Sciences, 17 July 2016.http://researcharchive.calacademy.org/research/entomology/Entomology_Resources/Hymenoptera/sphecidae/Family_group_names_and_classification.pdf

Useful websites:

http://www.antweb.org/
http://bugguide.net/
http://www.discoverlife.org/
http://evanioidea.info/

Mischocyttarus flavitarsis ♂

Bees of Southern Oregon (2016)

noreply@blogger.com (Travis Owen) — Wed, 09 Nov 2016 15:57:00 +0000

Hoplitis albifrons ♂

When most people hear the word bee, images of honeybees (Apis mellifera, Apidae) and the associated tales of declines come to mind. However, bees are much more diverse and interesting than the honeybee leads one to believe. There are at least thirty-five hundred species of native bees in America north of Mexico alone, and over twenty-thousand described species worldwide (probably many more). They live diverse lifestyles, but unlike the honeybee most live solitary lives. A similar variance can be seen in wasps, particularly predatory wasps which bees evolved from.

Xylocopa tabaniformis

Bees, wasps, ants, and sawflies make up the taxonomic order Hymenoptera, which literally means membrane-winged. Hymenoptera is derived from the Ancient Greek word humenópteros (ὑμενόπτερος), from ὑμήν (humḗn, membrane) and πτερόν ‎(pterón, wing). The most primitive members of the Hymenoptera are the sawflies (Symphyta, includes horntails and wood wasps) which lack the constricted waist seen in the other clades. True wasps likely evolved from sawflies, and bees evolved from predatory wasps (Crabronidae sensu lato) whose modern descendants often look similar to and are mistaken for true bees. The primary difference between wasps and bees is that bees have branched hairs on their bodies, believed to be an adaptation for carrying pollen, while wasps have simple hairs. While there is some truth in saying that bees are just fuzzy wasps, there are bees that are nearly hairless (i.e. Nomadinae and Hylaeinae) and wasps that are considerably hairy (i.e. Scoliidae).

Andrena sp.

Another difference between wasps and bees, and this is a very large generalization, is their diets. Wasps, either predatory or parasitoid, are for all intents and purposes carnivorous (at least in the larval stage) while bees are essentially vegan, feeding mostly on nectar and pollen. There are exceptions, for instance there are plant feeding wasps such as the gall forming wasps (i.e. Cynipidae) and bees that will occasionally be carnivorous such as when honeybees cannibalize their own larvae and eggs when other food sources are scarce. Also there are many adult wasps which drink flower nectar and others which do not eat at all in the adult stage. Ants, like bees, also likely evolved from predatory wasps or wasp-like ancestors, though unlike the other groups are entirely social (always live in colonies) and in general have a very wide spectrum of diets. Despite the name, velvet ants (or "cow killers," Mutillidae) are predatory wasps related so the social wasps (Vespoidea), females are wingless but males are winged and seldom seen.

Andrena pertristis ♀

Most bees, many modern wasps, sometimes sawflies, and occasionally ants visit flowers for their nectar and sometimes pollen, and are now considered to be important if not the most important pollinators. However, the earliest hymenopterans (Triassic, >250 million years ago) were probably plant tissue feeders as most sawfly larvae are today. The first flowering plants appeared in the early Cretaceous (~125 million years ago), and only produced pollen, and were wind pollinated. Nectar, or at least nectar-like sugary secretions, appeared some time around the late Cretaceous. Sugars were already present in plant tissues (there is no better example than the sugar cane plant, Saccharum sp.) and at some point in their evolution the plants began excreting the sugars through specialized organs called nectaries. It is easy to imagine an insect that once bit into the tissue of a plant for the sugar content to become gradually persuaded to visit the nectary instead, which would be mutually beneficial as it is less work for the insect and less damaging to the plant. Plants have used this strategy for other purposes than enticing pollinators. Bracken ferns (Pteridium spp.) produce a nectar-like substance from specialized glands (dubbed extra-floral nectaries) to entice ants which protect the tender new growth from being eaten from herbivorous invertebrates.

Apis mellifera in Ranunculus repens, an odd sight since pollen of Ranunculus is toxic to honeybee larvae. Specialists of Ranunculus pollen (i.e. Chelostoma florisomne, Osmiini) are able to metabolize the pollen just fine. Both A. mellifera and R. repens are of European origin, yet did not evolve together.

My own observations of the bees in my area and the observations of countless others have shown that bees of different types have very different preferences when it comes to flower types. Floral forms, to a great extent, dictate which types of bees (and other pollinators) can forage based on their body shape and tongue (proboscis) length. This is known as [the theory of] pollination syndromes. For example, large carpenter bees (Xylocopa sp.) have relatively long tongues and large bodies, while small carpenter bees (Ceratina sp.) have short tongues but their small size enables them to crawl into long floral tubes to reach nectar that may be inaccessible to the larger bees if it is deeper than their tongues can reach. The constituents of nectar and pollen vary considerably between plant species and the conditions in which they grow. This variation in nectar is also likely to contribute to which bees visit which flowers, though this is an understudied topic. Nonnative weeds and garden plants present a unique opportunity to study the subjects of pollination syndromes and bee species-specific preferences, as the bees have never encountered the exotic flowers before they were introduced.

Bombus vandykei ♂ and Ceratina sp. ♀

I work for a commercial beekeeper, and honeybees play an extremely important role in my life as they literally allow me to feed my family (by allowing me to earn a paycheck). For many others, honeybees are seen as the most important pollinator. In reality, honeybees are a commodity and are necessary to our food system primarily because we have plowed over the habitat of native bees and their native host flowers to place industrial scale farms and car-dependent low-density suburban wastelands that are inept at supporting much biodiversity.

Melissodes sp. ♂ (August)

Excuse me while I step up on my soap box, I promise it is restricted to this single paragraph. We need to provide habitat and flowers, preferably native flowers (or at least well behaved nonnatives) in order to promote native insects. By increasing in number, native bees not only pollinate more plants (including some that honeybees are unable or unwilling to pollinate) but serve as a food source for birds and other wildlife. Native bees have also been shown to improve the efficiency of honeybees on fruit or seed crops by forcing them to switch between crop rows more often. Most native bees are not aggressive and will more likely flee than sting, and some are too small to sting through human skin anyway. Besides all those good things, native bees are beautiful and diverse and I would like to see more in my garden and the world that my daughters are growing up in. Less Euonymus or Arborvitae, which support little to no biodiversity, more Madia and Horkelia, which support abundant biodiversity.

Below I have compiled all of the bees that I have photographed this year along with some of my observations of the species. They have been organized by their taxonomic classification (in other words, by scientific classification) for the sake of cleanliness. When I wrote about bees last year, I omitted honeybees from my report and focused solely on native bees that I had encountered. This year I have chosen to include them, mostly because I have been working with them intensely since April (Old Sol Bees). The bees in this report do not represent all of the bees in my area, just the ones that I was able to photograph. Please, enjoy:

Family Andrenidae

The Andrenidae represents one of the largest bee families (~3,000 species), and some of the most often encountered native bees. Many are small, much smaller than honeybees that is, though similar coloring would lead many lay observers to assume they are honeybees. This family includes the world's smallest bees, Perdita spp., which look nothing like their larger honeybee cousins and would most likely be mistaken for flies (Diptera). They are ground nesting bees which excavate their own branched tunnels two to three inches deep with a cell at the end of each branch (occasionally eight cells per burrow) with an egg laid atop a ball of pollen and nectar. They are solitary bees, though their nests may be in close proximity leading the uninitiated to believe they are eusocial and have colonies like honeybees. They often make their nests in sparsely vegetated areas, but are also known to nest in lawns where soil is showing. However, their small size and weak stinger equates to little to no threat to bare footed children.

Subfamily Andreninae

There are approximately fifteen hundred species in this subfamily, around 480 north of Mexico. The majority are in the genus Andrena, one of the most commonly encountered bee genuses in America, though many could be mistaken for honeybees to the unfamiliar.

Andrena pertristis ♀ on Hypochaeris radicata (May)

Andrena pertristis was one of the larger Andrena I have encountered, but still smaller than the average honeybee forager. It was foraging on the nonnative and noxious weed hairy catsear (Hypochaeris radicata), often mistaken for a dandelion (Taraxacum spp.), though they grow taller and bloom into summer and fall while the dandelions in my area bloom in late winter and spring. Catsear is perennial, grows from a thick and deep taproot, and seed is distributed by wind and birds. If plants are pulled, the taproot breaks and new crowns are formed. Any fragment of taproot forms a new plant, a highly successful species. Myriads of native bees flock to these flowers when in bloom.

Andrena pertristis ♀ on Hypochaeris radicata

Andrena sp. ♂ stealing nectar from Arctostaphylos viscida (March)

While all bees can fairly be called pollinators, sometimes they exhibit behavior that does not aid in pollination. Such is the case when the male Andrena steal nectar from the side of the manzanita (Arctostaphylos viscida) flower. Manzanita is in the heath family (Ericaceae) which includes blueberries (Vaccinium spp.), heather (Erica spp.), azalea (Rhododendron spp.), madrone (Arbutus spp.) and many others. The flowers of all of these require sonication, or buzz pollination, to release the pollen from the anthers. Andrenid bees are not typically capable of this, and it has been my observation that the andrenid bees in my area opt to rob the nectar instead. It is unclear whether they are the ones biting the holes or if it is done by other bees first.

Andrena sp. ♂ on plum blossoms (March)

Andrenid bees were the most numerous of all pollinators on my plum trees this year. When I would approach a tree a cloud of tiny bees would burst from the side I was on and move to the other side.

Andrena sp. on plum blossoms (March)

Unfortunately, the characteristics which define species of Andrena are minute and require a specimen to study or very detailed macro photos of various angles for a positive identification. My photos are not nearly detailed enough for this, something to strive for next year.

Andrena sp. ♀ on male Quercus kelloggii catkins (April)

Last year I had observed from a distance while pollinators visited the male catkins of the California black oak (Quercus kelloggii) but had not seen them close enough to determine what type of pollinator they were (hymenopteran or dipteran). Oak trees often bloom at the crown, so viewing pollinators visiting the typically wind-pollinated catkins is not typically practical. However, the oaks and many other spring flowers bloomed prolifically this year, and there were oak catkins within view enabling closer observation. Many Andrena appear to be generalists as in they visit a wide range of flowers across many families. It is doubtful whether pollinators contribute to the reproduction of oak trees since the female flowers, what will eventually grow to become acorns, offer no incentives for pollinators to go to them and are not located in the same area on the branches as are the catkins.

Subfamily Panurginae

This subfamily contains the worlds smallest bees in the genus Perdita. Of the thirty-five genera, many are specialists which collect pollen from a single plant species thus efforts to conserve species require the conservation of the host plant. Generally speaking, panurgine bees are less hairy then their Andreninae cousins, though their lifestyles are otherwise very similar.

Tribe Calliopsini

Calliopsis (Nomadopsis) ♀ on Horkelia daucifolia (June)

Species of Calliopsis are oligolectic, meaning they visit only a narrow range of flowers for pollen. I found a male and a female perched on flowers of Horkelia daucifolia (Rosaceae), a California and Southern Oregon endemic which is sometimes associated with serpentine. Some species of Calliopsis are specialists of a single plant species, though to my knowledge none are specialists of Horkelia spp., which appear to be generalist flowers (attracting many types of pollinators).

Calliopsis (Nomadopsis) ♂ on Horkelia daucifolia (June)

The male Calliopsis is identified by the lack of scopal hairs on the hind legs, which the female uses to collect pollen. Most bees (except for bees in the Apidae) have patches of thicker hair either on their hind legs or under the abdomen (as in Megachile spp.) which are used to hold pollen. Apidae bees have corbicula, similar to scopae except they form a sort of pouch sometimes referred to as a pollen basket. Males of any species of bee, as well as the kleptoparasitic bees (which do not collect pollen, but instead steal it from the nests of other bees), lack both scopae and corbicula.

Family Apidae

Within the family Apidae are the most common and familiar bees in America, including honeybees and bumblebees, both in the subfamily Apinae. Carpenter bees (subfamily Xylocopinae) are another familiar member of the family. The Apidae is the largest family in the superfamily Apoidea (all true bees) with >5700 known species. While the most well known members are eusocial, the majority live solitary lives. The family is composed of many unique and fascinating species, a few of which I will summarize here. Orchid bees (tribe Euglossini, mostly tropical) are an occasionally eusocial group, males of which collect oils produced from orchids (and plants in a few other plant families) to attract females, thus pollinating them. Cuckoo bees (subfamily Nomadinae) do not build their own nests, lack pollen carrying apparati, and lay their eggs in the nests of other solitary bees. Stingless bees (tribe Meliponini, tropical and subtropical) are small eusocial bees closely related to honeybees that are reared for honey and pollination in tropical parts of the world (not in the United States). Others include the digger bees and long horned bees, a group of fast flying solitary bees.

Subfamily Apinae

The subfamily Apinae includes all the eusocial species and the most familiar bees to most. Although honeybees (Apis spp.), bumblebees (Bombus spp.) and stingless bees (tribe Meliponini, many genera) all exhibit eusocial behavior, their methods of communication among nest mates and nest construction vary widely (even within genera) thus it is believed the social behavior developed independently in each group. This subfamily also includes all the solitary ground nesting bees of the Apidae.

Tribe Apini

The tribe Apini includes the single genus Apis, the honeybees. There are around twelve accepted species around the world, including the European honeybee (A. mellifera, which itself has around twenty six subspecies), the Asian honeybee (A. cerana), and the giant Southeast Asian honeybee (A. dorsata) which builds a large single wax comb in an exposed site like a branch or off the side of a cliff. Honeybees are probably the most evolutionarily advanced and complex of all bees, something I will attempt to explain in the following paragraphs. As mentioned earlier, many of my job duties involve working with honeybees (A. mellifera) in a variety of ways, so I should hopefully be able to adequately describe some facets of their complexity in a coherent manner. Though for me to think I can summarize a species whose literature can fill a modest library is absurd, I suggest that anyone interested in learning more about keeping honeybees should refer to the books Natural Beekeeping by Ross Conrad (Chelsea Green Publishing) or The Backyard Beekeeper by Kim Flottum (Quarry Books).

Apis mellifera queen ♀

One of my job duties for much of the year had been to capture queens to sell. This entails ascertaining whether or not a nucleus colony (a small hive created for the purpose of queen rearing, in our case) is queenright, has eggs present (indicative of the presence of a mated queen), locating said queen, capturing her barehanded, and placing her in a cage to be sent to a customer. After a queen is captured, and a hive becomes queenless, a new queen cell is added rather than letting them raise their own. My employer has been grafting honeybee larvae for about two decades, a process that allows control over at least some of the genetics while simultaneously creating as many queens as possible. Queen cells are placed (über carefully!) into the now queenless units where they will be raised by the adopted hive and, once fully mated (laying fertilized eggs), will be captured and the process repeated.

Apis mellifera queen (I swear I'm not squishing her! We capture and hold queens by the thorax, never the wings or abdomen, before placing them in a cage and given attendants to feed and groom them. We try to place them in the cage quickly since holding is certainly stressful, but if done right they will remain calm. Also, avoid breathing on them!)

Queens are created when a bee larvae feed exclusively on a diet of royal jelly, a protein rich substance excreted from the heads of nurse bees. All honeybees are fed royal jelly when they first hatch but only the new queens feed solely on the substance. In around sixteen days from the time the egg was laid by the previous queen, a newly pupated queen emerges, but she is a virgin. If there are other virgin queens present they will battle to the death, often by stinging. Queen bees, like wasps, have stingers that are not barbed like most bees. New queens will make a series of mating flights, mate with dozens of male honeybees (drones) and store all of their sperm in a special receptacle called a spermatheca. Once they are mated, they will never leave the nest again (unless they swarm). A mature queen is a workhorse, laying an estimated two thousand eggs a day in a strong colony.

Apis mellifera worker ♀ on Rubus armeniacus (Rosaceae)

Foragers are the most familiar of the honeybee castes. All forager honeybees are females, and all females can lay eggs, but they are only capable of laying males (drones) which will be explained in greater detail below. Queen honeybees suppress the tendency of worker bees from laying eggs with pheromones. Queen honeybees produce over two dozen distinct pheromones, the functions of some are as yet a mystery. Worker bees produce several pheromones themselves, which is a major part of their method of communicating with nest mates. They also communicate by touching one another, possibly taste as well, and by vibration. One of the most sophisticated of their communicatory expressions is known as the waggle dance. Dependant on the angle the dance is performed in relation to the sun, foragers are able to direct other foragers to choice sources of nectar or pollen.

Apis mellifera worker ♀

Newly hatched larvae of any caste are given royal jelly in the beginning, then if they are to be a worker or a drone they are switched to a diet of honey and pollen for six days. After pupating for about a dozen days under a wax cap, they chew their way out. The fuzzy new honeybees begin their lives as laborers by cleaning cells, then becoming nurse bees, then comb builders, honey/pollen packers, queen attendants, guard bees, then finally they retire as foragers. The order isn't rigid, sometimes they skip steps or skip around, occasionally reverting to earlier tasks. Most live out their lives foraging, a physically taxing occupation for the small insects. Honeybees have been known to forage two to five miles (radius) away from their hive. In the busiest parts of the season (for a honeybee, this is probably spring and summer when there are the most flowers), a worker may live for two weeks. In the winter when they are not flying, a worker may live for months.

Apis mellifera stinger

Like the predatory wasps, bees have stingers and venom. Unlike wasps, bees have barbed stingers which are torn out when they sting leaving a pulsing sack of venom, and some entrails, with the stinger. With the exception of queen honeybees, the bee dies after stinging. The honeybee stinger has multiple parts. The pointy part is made up of three parts, a thin channeled shaft (stylet) and two serrated parts (lancets) which glide up and down the stylet and drive the stinger into the victim. Meanwhile, two muscles continuously pump venom through the channel between the stylet and the lancets. In my experience, the sting of a honeybee is more painful than that of a yellowjacket wasp (Vespula spp.), but this is extremely variable and I suspect it is dependant on where the sting occurs on the body and the duration the honeybee stinger remains in the flesh (though I try to remove it as fast as I can).

Many of the more notorious hymenopterans (yellowjackets, hornets, Africanized bees) are preceded by their reputation as being aggressive. Fear of being stung has a very real effect on our perception of the order Hymenoptera, and for good reason. For a very small percent of the population (~5%) being stung just once can be life threatening. For the rest of us being stung is a painful warning and will have symptoms that go away. Thanks to the research of entomologist Justin Schmidt we now have a better understanding of insect stings, and the evolutionary significance of the fear they induce in us, which as it turns out is very important to their survival. Solitary hymenopterans, the bees and wasps that do not form colonies, have stings that, although occasionally extremely painful, do little actual harm. Eusocial hymenopterans, such as honeybees, bumblebees, and social vespid wasps, have venom that causes real harm by bursting cell membranes and destroying nerve tissue, among other things. This is significant, the social hymenopterans have nests full of valuable food stores and brood worth protecting while solitary hymenopterans, however, can abandon their small nest and create a new one elsewhere. Bees and wasps rely on the fear induced by the pain of their sting even though it is essentially a lie with the solitary species.

Apis mellifera drone (♂) perched on a daylily (Hemerocallis sp.) but you are unlikely to see a drone visiting flowers, because this photo was set up. I brought a drone home from work to show my three year old daughter (she loved it!), then released him the following day. He stumbled around stupidly for a few minutes in the daylily before clumsily falling off. Not one of nature's more graceful subjects.

Male honeybees are called drones, and their sole purpose is to mate. Drones, as is the case with all Hymenoptera, are born of unfertilized eggs. In other words, they have no father, and when their family tree is mapped out the resulting pattern that emerges is the Fibonacci sequence. One drone has one parent (a queen), two grandparents, three great grandparents, five great great grandparents, eight great great great grandparents, et cetera, ad infinitum. Queen bees can fertilize eggs at will by having full control of the spermatheca (sperm-storing receptacle) and ovary. A drone's life in the hive is that of luxury and indolence, until the time comes when the drones are forced out of the hive to seek a queen (by her pheromones), mate, and die. Unlike their sisters, drones lack a stinger but have a penis, but it is broken off at the time of mating. They typically bleed hemolymph and die following sex. Remember this next time you are having a bad day.

Following are a few examples of some typical honeybee apiaries, or bee yards. Typically there is a limited number of hives an area of land can support, based on the amount of resources available and if there are other apiaries nearby (though beekeepers tend to communicate on these things). Like other nonnative agricultural animals, honeybees do have an impact on the native ecology, though the full effects are not fully known. Unlike cows which can be confined to an area with a fence, bees cannot be fenced and thus directly compete with native pollinators. My personal observations tend to suggest that while honeybees may be creating more competition, the larger threat to wild bee populations is most likely loss of habitat and lack of native wildflowers in heavily human-populated areas. Some of the apiaries I have seen were located in land with very few invasive plant species and were able to support a wide variety of native bees despite the forty to seventy honeybee hives in the area.

Tribe Bombini

The bumblebees may be the second most recognizable group of bees after honeybees. The tribe contains the single genus, Bombus (46 species north of Mexico, ~260 worldwide). There are, however, species of bumblebee brood parasitoids that do not collect pollen and take over the established nests of pollen collecting species (by way of a coup d'etat of sorts, and mimicry) and have occasionally been considered a separate genus, Psythirus. Besides the parasitoids, the genus Bombus is a eusocial species, but unlike honeybees which have perennial colonies, have annual colonies. In the spring, a new queen bumblebee seeks a suitable nest site, often in old rodent tunnels or other protected spaces (including gaps in walls, discarded mattresses, etc.), and builds a small wax urn-like vessel for storing nectar. Once enough resources are collected she lays a few eggs to raise a few workers. When enough workers have reached adulthood she ceases to forage and focuses solely on egg laying. Depending on the species, males and new queen are produced some time around late summer or fall. Males and new queens mate, males die, and the newly mated queens either return to the nest temporarily or seek a winter hideout (hibernacula). The current years colony along with the old queen die, and the new queens start the process again the following year.

Bombus vosnesenskii ♀ foraging on Lathyrus latifolius

Bumblebees, along with a few other local genera (i.e. Habropoda: Anthophorini), are able to perform sonication, or buzz pollination. This is the process of vibrating certain flight muscles while gripping onto flowers in order to release the pollen from the anthers. This sort of movement is required for many plants including those in the Solanaceae (nightshades: tomatoes, potatoes, eggplant, etc.), Ericaceae (blueberries, cranberries, manzanita), and a variety of genera in other families (i.e. Dodecatheon: Primulaceae). Honeybees are inefficient or unable to pollinate many of these plants, thus the importance of alternative pollinators.

Bombus vosnesenskii ♂ on Hyssopus officinalis (June)

Bumblebees are not as sophisticated as honeybees when it comes to communication, but this has major benefits for people. This is because bumblebees do not communicate where good floral resources are like honeybees (the waggle dance). This may not seem beneficial right away, but in the case of greenhouse pollination it ensures pollination of the target crop. Honeybees, being the fickle little girls they are, will forsake one plant altogether if they find something more appealing. This could result in the target crop being ignored completely, which is an obvious drawback. In the case of greenhouse tomatoes, which require cross pollination, yet are not well liked by honeybees, bumblebees do a fine job. Many people are unaware that bumblebee colonies of a limited few species are commercially available, but once again they are "single use" as the colony will die at the end of the season. Being that there are a limited number of species that are reared commercially, it is of high importance to only use a species that is native to your area lest one should introduce yet another nonnative species to what might have been healthy ecosystems.

Bombus vandykei ♂ on Allium sphaerocephalon (June). Males like this one are often found motionless on flowers early in the morning, offering excellent opportunities for the photographer.

A few more comparisons to the honeybee, the first being that bumblebees do not build hexagonal cells arranged on flat vertical combs as do honeybees, and the second is that they do not make honey (despite collecting nectar). Bumblebees are essentially a ground nesting genus as opposed to honeybees which prefer nest sites with a bit of elevation off the ground), and their nests are constructed by a nonuniform jumble of wax pots filled with nectar, some pollen, and brood. Since they don't form perennial colonies, they have no need to make honey (which is essentially created for the purpose of lasting a long time, as evident by unspoiled three thousand year old honey found in ancient Egyptian tombs). Despite these comparatively more primitive attributes, bumblebees were for thousands of years the dominant eusocial bees in the Americas before the introduction of the honeybee in the time of the European colonization.

Bombus huntii ♀ and Echinacea purpurea (July)

I have observed a progression of species of bumblebees in my area. The first to appear are queens of Bombus vosnesenskii and B. vandykei. In about a months time their workers become apparent, and the bright yellow males of B. vandykei are conspicuous visitors to the garden. Later in the season, as B. vandykei become less numerous, B. huntii appear, though I have observed fewer of them than the previous two. I have seen a few other species, including B. griseocollis and a few other species whose identities remain a mystery. Bumblebees are not too difficult for the lay observer to identify. They differ from the large carpenter bees (Xylocopa spp.) by their hairy abdomens, whereas the abdomens of the carpenter bees are shiny and hairless. Bumblebees are identified by the bands of colors on their bodies, often by the location of particular colors on the abdomen. For instance, B. vosnesenskii females differs from B. vandykei only by the the location of a single yellow band on one abdominal segment rather than the one next to it (yellow band on abdominal segment T4 is B. vosnesenskii, T3 is B. vandykei). In other instances identification is much less subtle and more readily apparent.

Bombus huntii ♀ and Echinacea purpurea (July)

Bumblebees are generalist pollinators, like most bees, which means they will visit a wide range of flowers. In my garden they are quite fond of the masses of Echinacea I have grown from seed. Other plants they seem to be attracted to are, in no particular order, onions (Allium spp., including ornamental types), winter heath (Erica spp.), true hyssop (Hyssopus officinalis), manzanita (Arctostaphylos spp.), Anchusa spp., and Pacific hounds tongue (Cynoglossum grande), to name a few. Bumblebees are also important pollinators of many native orchids including Spiranthes spp., Piperia spp, and Goodyera spp. among others. In the case of orchids, the bumblebee is occasionally duped into visiting the flower which has no floral rewards, while in other cases some species of orchids actually produce nectar as an incentive to pollinators.

Tribe Emphorini

This is a rather poorly represented tribe of bees in the United States, with just thirty-one species in four genera north of Mexico, one hundred eighteen species in ten genera worldwide. They are known collectively as chimney bees for their occasional nest entrance featuring a turret made of mud. Most are pollen specialists of a relatively confined list of plant families, predominantly the mallows (Malvaceae) but also plants in the families Asteraceae, Cactaceae, Convolvulaceae, and Onagraceae.

Diadasia sp. ♂ inside Sidalcea glaucescens (May)

Like others in the tribe, Diadasia spp. are specialists. I have only observed this genus twice, but both times were in the waxy checkerbloom, Sidalcea glaucescens (Malvaceae), a common wildflower in my area. S. glaucescens has flowers that close at night, and this male was found inside this flower early in the morning. The fact that it was too cold for the bee to move yet the flower was open suggests the flowers are sensitive to light, but this is just conjecture.

Tribe Eucerini

This tribe is comprised of thirty six genera with around seven hundred fifty species worldwide, and two hundred twelve species in fourteen genera north of Mexico. They are known as long-horned bees because the males of most species have very long antennae. They are all ground nesting bees, occasionally building horizontal burrows in exposed banks. Like many of the ground nesting bees, females line the cells with a wax-like substance. Some species nest communally (multiple laying females sharing a nest) but most are completely solitary.

Melissodes sp. ♀ on Madia elegans (July)

Melissodes are specialists of flowers in the Asteraceae, or sunflower family, while the species in my area appears to specialize on the native Madia elegans. The flowers begin blooming in mid simmer and continue until mid fall. They open in the evening and remain open until the sun comes out. The bees work the flowers in the cool mornings before the flowers close, then disappear for the remainder of the day. July is one of the hottest months of the year here, often in the 90° F range but occasionally hotter.

Melissodes sp. ♂ on Madia elegans (July)

These are very fast flying bees, and very difficult to sneak up on. Males sometimes fight over females, and males also attempt to mate while females are foraging. I witnessed a mating event in which a female visiting a flower was suddenly mounted by a male, and they struggled briefly before the female was able to fight him off. I have no idea if the mating was successful, it was less than thirty seconds.

Subfamily Xylocopinae

The Xylocopinae is the third subfamily in the Apidae (following the Apinae and Nomadinae), and represents the carpenter bees. There are four tribes in the Xylocopinae (Allodapini, Ceratinini, Manueliini, and Xylocopini), only two of which are represented north of Mexico. The tribe Allodapini is represented by around fifteen genera, mostly in Southeast Asia, sub-Saharan Africa, and Australasia. They often form communal colonies where there are multiple laying females caring for brood, but what really makes this tribe strange is the larvae are entirely mobile within the nest, and must compete for food. The tribe Manueliini is represented by a single genus with a handful of species, and are only known from Chile and Argentina. The tribes Ceratinini and Xylocopini are a bit more familiar, and are described below.

Tribe Ceratinini

The single genus, Ceratina, represents this tribe. They are small bees that nest in the pith of broken stems of woody plants (such as Rubus spp.) or occasionally wood itself though I would assume it would be very soft wood. They are mostly solitary, though there have been reports of some species exhibiting eusocial behavior where one bee forages while the other lays eggs and guards the nest. They often appear black though they are typically dark metallic green or blue. Interestingly, females do not collect very much pollen on their sparsely hairy bodies, but rather eat it and store it in their crop until they return to their nest. There are other some other bees that collect pollen in this way (i.e. Hylaeus spp.: Colletidae, which I photographed here last year), mostly small bees.

Ceratina sp. on Hypochaeris radicata (May)

There are about twenty one species of Ceratina in the United States, mostly found in non-desert habitats. Most American species are solitary and nest in broken pithy stems. The females create a linear row of brood cells, starting from the deepest, and separate the cells with the chewed up pith and their own saliva. When the nest is complete, the female will stand guard at the entrance of the nest, dying in winter or late fall (first hard frost), and remain in place to block the entrance from intruders. Larvae pupate in the fall and remain in the nest in diapause as adults through winter until they emerge in the following spring.

Ceratina sp. on Hypochaeris radicata (May)

Ceratina appear to be quite fond of plants in the Asteraceae, though this could be that many composites have relatively accessible nectar and pollen. Ceratina are generalists, and readily accept foreign plants such as Hypochaeris and Echinacea as pollen sources. In general brood are more susceptible to different sources of pollen than adults are of nectar because pollen has so very much to do with development. Pollen is the primary source of protein for bees, and protein content varies greatly between plant genera. Wind pollinated plants have pollen that is practically devoid of protein, suggesting that protein in pollen was an adaptation to entice pollinators. Studies where pollen specialist bees were fed different pollen sources resulted in larvae that didn't survive to pupation, or were greatly reduced in size as adults. Generalist species like Ceratina have clearly adapted to be able to metabolize a wide variety of pollen types and complex sugars, and it is no surprise the genus is found in every continent except Antarctica.

Tribe Xylocopini

The most iconic members of the subfamily Xylocopinae, Xylocopa is the sole genus of the tribe Xylocopini. These large conspicuous bees make nests in living and dead wood or woody plant stems (i.e. bamboo, Agave spp., Yucca spp.) with their strong mandibles. Nests are usually linear, occasionally with brood cells coming off a main tunnel. Most are solitary, though a few are primitively eusocial as females of overlapping generations share a nest. Cell walls are lined with sawdust, and species of desert habitats line cells with a wax like substance to prevent desiccation. The eggs of some Xylocopa are the largest insect eggs known, some over a half-inch long! Females lay few eggs in their lifetime, often less than eight, and relative to other bees invest greater maternal care in rearing them.

Xylocopa tabaniformis visiting Salvia sclarea (July)

Males of Xylocopa are often seen lurking around flower patches they have deemed good places to pick up hot dates, and will aggressively go after any intruder that happens by, even flies (Diptera), and sometimes people. There is nothing to worry about though, like most male hymenopterans, they have no stinger. When a female comes along, they will attempt to mount her and remain attached while she attempts to continue foraging.

Xylocopa tabaniformis visiting Salvia sclarea (July)

Large carpenter bees were one of the only species to show interest in the clary sage (Salvia sclarea), along with carder bees (Anthidium manicatum), a few European paper wasps (Polistes dominula), and a hummingbird (Selasphorus rufus). They were, however, highly interested in this plant and there were at least a handful of them working the clary sage at any given moment while they were still active. Although clary sage is native to the Mediterranean, its reproductive structures are a good fit for the large carpenter bees. A few years ago I grew Salvia farinacea (before the voles killed all of my plants) and the carpenter bees were similarly attracted to those flowers as well. Monarda fistulosa is another plant in my garden that is attractive to Xylocopa tabaniformis.

Family Halictidae

Halictid bees are only second to Apidae in terms of numbers, and include some species that are readily recognized by some gardeners and naturalists. They are collectively known as sweat bees, though this is misleading as only some members of the family land on people and are attracted by sweat (some species in the tribe Halictini). Certain flies in the family Syrphidae (Diptera) are also known as sweat bees in some regions since some species are attracted to sweat. Halictid bees are typically ground nesting bees, some prefer clay soil while others prefer sandy soil at the edges of streams. Some have a single generation per year, but most probably have multiple overlapping generations. Most are fairly considered to be solitary, though some become eusocial as more generations emerge from pupation and share the nests with their mothers. Others are strange in that, within a single species, may be solitary in one region while eusocial in another. This is thought to be determined by climate and how many resources are available. Others may simply be communal, with females sharing a single nest entrance but provisioning their own brood cells.

Subfamily Halictinae

This subfamily contains the most commonly encountered genera, including the small metallic green bees in the genera Agapostemon and Augochlora (Augochlorini). Some are even kleptoparasitic, the genus Sphecodes isn't commonly encountered and resembles a wasp (black mostly hairless body with a red abdomen), and it lays its eggs in the nests of other Halictinae.

Tribe Halictini

The tribe Halictini contains at least 2,800 species in twenty two genera in the world, three hundred eighty species in five genera north of Mexico.

Agapostemon sp. ♀ on Hypochaeris radicata (May)

Agapostemon spp. are one of the easiest of the green metallic bees to identify by the lay observer. The head and thorax are metallic green and the abdomen has black and white stripes. These are typically small bees that visit a wide range of flowers, mostly Asteraceae in my observations. Although they are sometimes known as greed sweat bees, Agapostemon spp. are not attracted to sweat.

Agapostemon sp. ♀ on Hypochaeris radicata (May)

Agapostemon spp. are not eusocial, but some live communally. Occasionally a single female will guard the communal nest entrance, blocking access with her body, and only her head visible from above ground.

Halictus ligatus ♀ visiting Croton setigerus (July)

One of the most common native bees in my area is the sweat bee Halictus ligatus. This is a widespread species found from Canada to the Gulf of Mexico, and from the Atlantic to the Pacific. This species, like others in the genus, are true sweat bees because they are attracted to sweat. Halictus nest in flat ground in what may best be described as sandy loam. Tunnels may be simple or branched, and brood cells are lined with a waxy substance from a special organ at the tip of their abdomen called the Dufour's gland.

Halictus ligatus visiting Coreopsis (June)

Females begin foraging and creating solitary nests in the spring, but may develop into eusocial colonies. Eusociality ranges from having communal nest where overlapping generations remain in the nest to help care for younger generations to full hierarchical social systems with a single laying queen and worker caste. Dominant queens aggressively defend their role as the primary egg-layer (and the sole bee to pass on her genes), but depending on climatic conditions may have to contend with other laying queens in the nest. Favorable conditions this year will result in [relatively] large-bodied queens next spring, and if next year presents poor climatic conditions during the rearing of the first generation of workers they will be resultantly smaller in body size, thus easy to control by the dominant queen. The reverse scenario would lead to more laying workers, but unlike honeybees they would be entirely capable of mating and laying females. (In a honeybee hive, non-queen bees, known as laying workers, are only capable of laying drones, male honeybees, which have only one set of chromosomes from the mother.)

Halictus rubicundus ♀ foraging on Horkelia daucifolia (May)

Similar to Halictus ligatus, H. rubicundus is a somewhat abundant ground nesting species. Similar eusocial variability is found in this species, where cool climates encourage solitary nesting behavior, and warm climates encourage eusociality, and a mix inbetween.

Halictus rubicundus ♀ (May)

I witnessed this bee only once (that I am aware of), and it was one of the many pollinators visiting Horkelia daucifolia. The plant is a rose relative (family Rosaceae) with a fairly restricted distribution in Southern Oregon and Northern California.

Lasioglossum sisymbrii ♀ (January)

The genus Lasioglossum is very similar to Halictus in both morphological and variable eusociality. They are extremely common, but are easily missed due to the small size of many species. Like Halictus they are attracted to sweat, perhaps for nutrients in the salt needed for reproduction. Brood cells in underground burrows are lined with a glandular secretion of a mixture of chemicals called lactones, and some species line the entire nest with the waxy substance.

Lasioglossum sisymbrii ♀ (left) and Apis mellifera ♀ (right)

I discovered this sweat bee in mid January, apparently attracted to my beehive. Often our native bees are mistaken for honeybees by the untrained eye, but seeing both this Lasioglossum sisymbrii and Apis mellifera together there are some clearly recognizable differences. This was an odd sighting for another reason since it was so very early in the year. Mated females of both Halictus and Lasioglossum overwinter as adults, so this one could have been overwintering in or around the beehive, attracted to the warmth generated by the honeybees and the sugars that abound in its presence.

Lasioglossum sp. ♀ (Hemihalictus series) on Triteleia hendersonii (May)

This is one of the many small species of Lasioglossum that could be easily overlooked. The Hemihalictus series is a group of taxa that contains about nine subgenera of Lasioglossum with weak venation in the wings. They are short tongued bees, so often will be found on flowers with open and highly accessible floral rewards. This female was busy collecting the blue pollen from a yellow form of Triteleia hendersonii, a native wildflower that grows from a corm.

Lasioglossum sp. ♀ (Hemihalictus series) on Triteleia hendersonii (May)

The pollination ecology of many native bulbs, particularly in the genus Triteleia (and others in the Brodiaeoideae, formerly Themidaceae), is relatively unknown and obscure. Bees have been assumed the primary pollinators of this group of plants, but few documented observations are available. Small bees like the weak-veined Lasioglossum may be responsible for their reproductive fitness.

Family Megachilidae

This family is diverse and widespread, found in every continent except Antarctica, represented by over four thousand species in around eighty genera. North of Mexico, there are at least six hundred thirty species in eighteen genera. They are probably most distinguished by their namesake, the leafcutter bees (Megachile spp.) whose species use leaf pieces to partition and seal brood cells, but not all taxa use leaf bits. For example, other taxa include the mason bees (Chelostoma and Osmia spp.), carder bees (Anthidium spp.), and the megachilid cuckoo bees (Stelis and Coelioxys spp.). The binding characteristic among all megachilid bees (with the exception of males and the cuckoo bees) is that they carry pollen on scopae on the underside of their abdomen rather than on their legs. Pollen on females (since neither males or cuckoo bees collect pollen) is often conspicuous since the underside of their abdomen will be brightly colored yellow or orange, or whatever color the pollen is on the plant they are foraging on.

Subfamily Megachilinae

The subfamily Megachilinae is the largest within the Megachilidae with over seventy genera worldwide. The blue orchard mason bee (Osmia lignaria) and the blueberry bee (O. ribifloris) are the most economically significant and recognized of this group, primarily because they are sometimes reared commercially and sold across the United States, often where they are not native.

Tribe Anthidiini

Anthidium manicatum ♂ on Stachys byzantina

The European carder bee (Anthidium manicatum) is well introduced to the state of Oregon, where it is associated with Stachys byzantina (lambs ear), from which it gathers the hairs that cover the leaves to line its cells. They also use a variety of other materials in their nest construction, including plant resins and mud. Males are territorial and can be seen chasing off other males, or anything that flies nearby. I've even seen them go after and grapple with a bee fly (Bombylius sp.) trying to visit the Stachys flowers. Alas, these bees are fast and agile fliers which have the ability to hover and fly sideways, and have proven themselves a challenge for me to photograph. They are one of the few bees attracted by the flowers of clary sage (Salvia sclarea), and otherwise seem to have an affinity for plants in the Lamiaceae.

Tribe Megachilini

The most renowned members of the Megachilidae are the true leafcutter bees in the genus Megachile. To some they are considered to be pests because of the damage they can inflict on the leaves of some plants. As a rule, Megachile spp. use their mandibles to cut out semicircular sections of certain leaves to use in their nests. Nests are often in preexisting holes in wood, such as those from wood-boring beetles, and leaf tissue is used to partition cells and seal the nest entrance once the cells are full. Nests are linear, with some pollen, nectar, and a single egg in each cell.

Megachile sp. ♀ on Lathyrus latifolius (August)

I have in years past seen Megachile bees on only a few plants. The first was on Rudbeckia hirta, and the second sightings were on the perennial sweet pea, Lathyrus latifolius. They have a characteristic sweeping motion as they collect pollen, being as the scopae is located under the abdomen they must brush themselves over the anthers to collect the pollen. I regret not having a better photo, but just as I was about to photograph the bee in this photo a second time a [expletive] honeybee came and bullied the leafcutter bee away.

Tribe Osmiini

The Osmiini is best known for the genus Osmia, commonly known as mason bees. There are roughly eleven hundred species in seventeen genera worldwide with three hundred species in eight genera north of Mexico. Nest types and materials vary widely, some Osmia sp. have even been known to nest in empty snail shells. Nests can either be in preexisting holes in dead wood or in underground burrows, and anything from plant resin to leaf tissue to mud and pebbles is used to partition cells and seal nest entrances. Many specialize on one or a few plant families, some specialize on a single genus.

Ashmeadiella sp. on Coreopsis (June)

Many of the osmiine bees are pollen specialists of plants in the Asteraceae, while many genera in the Asteraceae are generalist flowers which have shallow and easily accessible floral rewards.

Hoplitis albifrons ♂ on Wyethia angustifolia (May)

Hoplitis build solitary nests with mud walls in a variety of locations depending on the species, from nests in the ground to aboveground nests in pithy stems of woody plants, holes in wood, or nests of mud dauber wasps (i.e. Sceliphron spp., Trypoxylon spp.). A few are even known to nest in old snail shells, and have been known to use rabbit dung to line the cells.

Hoplitis albifrons ♂ on Wyethia angustifolia (May)

Last year (2015) I observed bees within thirteen genera from five different families. Combined with this years (2016) observations, I have photographed a total of nineteen genera. This amounts to at least twenty-six species total, or twenty-four species of native bees (Apis mellifera and Anthidium manicatum are introduced). Perhaps I can inspire some of you to go watch some flowers, observe bees, and perhaps make some interesting observations of your own. Observe how different types of bees prefer different types of flowers, how they move differently, whether they are collecting pollen or nectar. A warning, it can be quite addicting and, in our current world, a source of optimism and hope.

Useful Websites:

http://bugguide.net/node/view/59
http://www.inaturalist.org/
http://www.discoverlife.org/
http://blogs.ethz.ch/osmiini/

Melissodes sp. ♂

Polistes aurifer

noreply@blogger.com (Travis Owen) — Tue, 08 Nov 2016 15:38:00 +0000

Polistes aurifer queen on her nest

While working in one of the apiaries recently, I had discovered a nest of the native Golden Paper Wasp, Polistes aurifer, underneath some empty bee equipment. There were two or three workers and the queen who was unwilling to leave. While I had my beekeeping veil on, I decided to get some photos. I question whether or not I even needed a veil since there were no signs of aggression. The workers simply fled while the poor queen was left with her brood.

Mischocyttarus flavitarsis

Although paper wasps and other social wasps have the reputation for being aggressive, this species (and perhaps the entire genus, ~19 species) and the related Mischocyttarus is relatively gentle and nonaggressive. One key difference I have observed is that the colonies of P. aurifer remain relatively small and may have up to 300 cells per nest and comparatively few individuals. I have personally not seen a nest larger than my fist. For comparison, honeybee colonies can have upwards of 100,000 cells and 50,000 individuals in a hive. Bald faced hornets (Dolichovespula maculata) and other aerial yellowjackets build nests the size of a large watermelon with 700+ individuals while true hornets (genera Vespa) build nests in similar proportions. Yellowjackets in the genus Vespula can have underground or otherwise sheltered colonies that attain massive size with over 15,000 cells and several thousand individuals. Dolichovespula, Vespa, and Vespula are much more numerous, defensive, and potentially aggressive than Mischocyttarus and Polistes.

Polistes aurifer (L) and P. dominula (R)

A common European invader to much of the United States, the European paper wasp (Polistes dominula, pictured above on the right) has nests that look nearly identical to the nests of P. aurifer except for occasionally attaining nest sizes similar to bald faced hornets (Dolichovespula maculata). This species is slightly more aggressive, and often shares habitat with P. aurifer (which in my case is under the eves of my roof). They can be told apart by their patterns, higher contrasting patterns of P. dominula while the abdomen of P. aurifer is mostly a dull yellow. I would also like to add the observation that the antennae differ in color, P. aurifer antennae fade to black at the ends while P. dominula antennae are black at the base. Markings on the thorax of Polistes dominula are also diagnostic.

Polistes aurifer ♂

Males lack stingers, like all other male Hymenoptera, and are produced towards the end of summer or the beginning of fall. They will mate with new females and die. Newly mated females will hibernate in a protected location through winter and build new nests in the spring, sometimes competing with other females over who will be the dominant layer. This appears to result in lost limbs of laying females, a consequence of retaining genetic superiority.

Polistes aurifer

Polistes aurifer nests are made of plant fibers and wood pulp, so in other words, paper. Wasps as we know them first appeared in the Jurassic and social wasps appearing in amber sometime around the Upper Cretaceous period. The main diet of Polistes and other vespid wasps (Vespidae) is comprised of chewed up caterpillars for their developing larvae and either nectar or honeydew (the excrement of aphids and other sap sucking insects) for adults. Bees, in contrast, procure protein for their young from pollen while vespid wasps in general attain protein from other insects and can fairly be called predatory. This trait can be very beneficial in a garden where caterpillars are a problem.

Polistes aurifer eggs

Eggs of Polistes aurifer are larger than honeybee eggs and more elliptical. Also note that the eggs of this species are towards the inner side of the cell, honeybees in contrast are always in the center of the bottom of the cell. It takes roughly 60 days for a mature P. aurifer to emerge, around 20 days each in the thee stages of development: egg, larvae, pupae. Honeybees, in contrast, take only 16 (queen), 22 (worker), or 24 (drone) days to emerge as adults.

Polistes aurifer

When I was watching this creature, I did not see an aggressive pest, but rather a mother guarding her young. When I first learned about this species, I thought twice about thoughtlessly spraying the nests in my eves or elsewhere. Had anyone been stung? No. I was just following the status quo and not considering my impact. By indiscriminately killing native wasps or other insects, I could be seriously disrupting the natural balance, and it just may manifest itself into a pest infestation in my garden or other unforeseen issues. Aside from all the pest control services these insects provide, they are also prey themselves to birds and likely other creatures in the food web. Why mess with something I do not fully understand, and why do it without fully considering the consequences? Why poison the native (and did I mention free?) pest control service providers? So I urge you to reconsider what you know and to ask yourself if you really know enough. I suspect as a species we do not know enough about the natural world as it is nor do we know the full consequences of our tactics. It's worth considering.

Polistes aurifer

Colchicum as a Pollinator Plant

noreply@blogger.com (Travis Owen) — Mon, 10 Oct 2016 13:30:00 +0000

Colchicum cilicicum with Apis mellifera.

Plants that captivate me are simultaneously uncommon in gardens, have unique life cycles, and are attractive to pollinators. One such group of plants that meet this criterion are in the genus Colchicum. Bees and some other pollinators have been captivated by these flowers, which grow from corms (though this is debated, their morphology is unique). Despite the colloquial name, "autumn crocus," some of the 100-150 species bloom in autumn, while many others bloom in winter and spring (and in the genus Crocus, there are many autumn and winter blooming species as well). The most commonly available species and hybrids bloom in the fall without their leaves, while the leaves and seed pods appear in the spring. They are a late source of pollen and nectar, and thus are very attractive to bees and other pollinators.

Colchicum cilicicum growing among Geranium macrorrhizum leaves

Colchicum are primarily bee-pollinated, though flies and perhaps certain wasps may play larger roles in pollination in some districts. The closely related genus Androcymbium has some species that are pollinated by rodents and perhaps birds in their native South Africa. Androcymbium has flowers that are small and held on small pedicels grouped in heads within green or petaloid bracts, lacking any resemblance to Colchicum as it is commonly known. The flowers of Colchicum sensu stricto lack above-ground stems and are therefore weak, held up only by the calyx, thus unfit for pollination by vertebrates. Androcymbium is found mostly in continental Africa with at least one Mediterranean species while Colchicum sensu stricto is native to Europe, North Africa, and Western to Central Asia.

Colchicum cilicicum leaves are pleated but not too large.

The leaves of some species appear in the spring, while others appear in the fall or in winter, sometimes while blooming or just after the flowers fade. Species which flower and leaf out in the winter tend to be native to milder, perhaps maritime, climates that do not get too cold in winter. The species and hybrids I grow are perfectly hardy, flowering in the autumn and showing little signs of growth until spring when the leaves and seed pods emerge. My garden is home not only to me and my garden but to deer, voles, wild turkeys, rabbits, tree squirrels, and red digger squirrels (among others) and the colchicums are ignored by all of these herbivores, both above and below ground, at least in part due to toxic alkaloids within the plants. Slugs are a minor nuisance, occasionally eating parts of the leaves, but never enough to cause concern.

Colchicum cilicicum with Apis mellifera

Why flower in autumn? There are a number of reasons why some species of Colchicum have adapted to flowering in the fall, winter, or spring. All geophytes (which possess at- or below ground storage organs; i.e. tubers, bulbs, etc.) have the advantage over non-geophytes of flowering outside of the typical growing season, and/or when conditions are not ideal for growth (too cold, dry, etc.) by using stored energy rather than waiting for optimal conditions. This results in less competition by other plants, and enticing (or forcing?) pollinators to visit their flowers. Large showy flowers in a part of the year that is otherwise scarce or devoid of other flowers is a huge advantage. This is apparent in my garden where multiple species of bees and flies visit the few specimens I grow. At any other point in the year, a single specimen of a given plant species is unlikely to attract pollinators, especially a diversity of pollinators, because pollinators in general are more interested in large groups of flowers. Autumn blooming species seem to be the exception to the rule.

Below are a few selected fall blooming species and hybrids from my own garden and abroad, with photos by Ian Young (Scotland), Gideon Pisanty (Israel), and myself (SW Oregon), with an emphasis on pollinators:

Colchicum stevenii with Andrena sp. (Photo by Gideon Pisanty, 2009)

Colchicum stevenii is a Mediterranean species native to Cyprus across Southern Turkey to Syria, Jordan and Israel. It grows on rocky slopes and dry banks and blooms from October to December, flowering beginning after the first fall showers. The leaves often appear with the flowers or sometimes as the flowers fade. Usually, only one or two flowers appear from each corm, but clumping individuals may produce bigger displays. Colchicum stevenii is a widespread species where it is native, though not oft grown in gardens. These photos by entomologist, researcher, and photographer Gideon Pisanty of Tel Aviv University show Colchicum stevenii in its native habitat in Israel being visited by native andrenid bees.

Colchicum stevenii with Andrena sp. (Photo by Gideon Pisanty, 2009)

Colchicum stevenii produces some nectar, but it is in small quantities and is dilute (low sugar content). To make up for this it produces pollen in good quantity, yet the species is polymorphic, having at least five distinct forms. Stigma length varies from short (below anthers), medium (level with anthers), and tall (higher than anthers) in addition to having both female (lacking anthers) and male (lacking stigma) forms. Different pollinator types visit each form. In one study (Dafni 1996) solitary bees and honeybees were observed to be most attracted to forms with short stigmas, while syrphid flies were most interested in flowers with long stigmas. Solitary bees were found to be more interested in the nectar than honeybees, and I would guess this to be due to the typically smaller body size of solitary bees than honeybees.

Colchicum 'Lilac wonder' with Halictus ligatus

A more accessible and commonly grown colchicum is the hybrid Colchicum 'Lilac Wonder'. It is an old hybrid dating from at least 1926, if not earlier, and speculated to be the progeny of C. giganteum and C. bornmuelleri (both occasionally considered synonyms of the variable C. speciosum), or possibly C. speciosum and C. bivonae (a species commonly used in breeding efforts). Whoever the parents are, this hybrid has very large flowers that can span five inches across when fully opened, and stand ten inches tall. This leaves it susceptible to mechanical damage from wind, rain, or animals (including children, like my three year old), leaving them laying flat like deflated balloons after the kids birthday party.

Colchicum 'Lilac wonder' with my hand for scale

Perhaps as a result of it being a hybrid, 'Lilac Wonder' seems to produce pollen in only small quantities which lessens its usefulness to pollinators. Despite this, it is still visited by a variety of bees. In my observation this is usually honeybees, but native bees occasionally visit these. I have not seen bumblebees visit the colchicums, though this is most likely because the Bombus species in my area are not active at this time of year with the exception of a few of next years queens.

Colchicum 'Lilac Wonder' with Toxomerus marginatus

Flower flies, such as syrphid flies, primarily feed on pollen and so are attracted to the anthers of Colchicum. Most of the floral visitors that visit the colchicums in my garden are interested in the pollen, while very few are interested in the nectar. I wonder if most species have poor quality nectar like C. stevenii, or if that is variable between species or climate. Note the lack of pollen on the anthers of C. 'Lilac Wonder' compared to the easily visible pollen in the photos of C. stevenii and C. cilicicum.

Colchicum speciosum

Another Colchicum I grow is C. speciosum, a commonly available species that is used frequently as a parent of new hybrids. For me, it is the first to bloom, but it does not seem to thrive in my USDA zone 7 climate. It does not seem to be as attractive to pollinators in my garden, either.

Colchicum speciosum with Eupeodes sp. (top) and Eristalis sp. (bottom) (Photo by Ian Young, 2016)

Halfway across the world, in the garden of Ian Young in maritime Eastern Scotland, Colchicum speciosum seems to perform much better. This could be climate, or it could be that the species is grown by a gardener with more skill than I. Flower flies in the family Syrphidae often have similar colors and markings to bees and wasps, possibly warding off some predators expecting a stinger which they do not possess. Most of these flies feed on pollen but they will also feed on easily accessible nectar. Colchicum has nectar that is only accessible to insects with a proboscis long enough to reach it, and most flies have short proboscises that are better suited to shallow flowers such as those found in the Apiaceae (carrot family). Pollen then, it would seem, is the primary reward to pollinators in Colchicum.

Colchicum speciosum leaves

Leaves of the many species can vary quite a bit in appearance. These are the leaves of Colchicum speciosum in my garden, compare to the leaves of C. cilicicum above. Some species, like C. montanum, have very thin strap-like leaves that are more comparable to the leaves of Muscari than C. speciosum or C. cilicicum.

Colchicum montanum with Eupeodes sp. (hoverfly) (Photo by Ian Young, 2006)

Colchicum montanum (syn. Merendera montana) is a European species native to the Central Pyrenees and Iberian peninsula from 900 to 2500m. It grows in grassland and rocky slopes, and is often grown in rock gardens and alpine houses (photos seen here by Ian Young in his bulb house). I speculate that flies are the primary pollinators, being a high altitude species, where flies are more reliable pollinators than bees (although both are present) in alpine and tundra environments. This would account for the comparatively equal length, yet short, reproductive parts.

Colchicum montanum with Eupeodes sp. (hoverfly) (Photo by Ian Young, 2006)

Colchicum montanum shares its environment with the Mediterranean pine vole (Microtus duodecimcostatus) which, despite the toxic alkaloids in the corm, eat the corms. This actually seems to enhance the fitness of C. montanum by spreading small corms about and reducing competition between individuals. The species seems to lack a seed dispersal method that separates plants, so the voles may be playing a role in inadvertently distributing the corms once established.

Colchicum (bivonae?) hybrid with Vespula vulgaris (Photo by Ian Young, 2004)

Ian Young grows a number of colchicums, species and hybrids, in his garden in Eastern Scotland. Photographed here is an old hybrid, at least two decades old, with a common yellowjacket, Vespula vulgaris. The name of this hybrid is long lost, although it bears a resemblance to Colchicum bivonae which has been used as a parent to many hybrids lending to them its petal tessellation.

Colchicum (bivonae?) hybrid with Vespula vulgaris (Photo by Ian Young, 2004)

While wasps are probably less frequently seen visiting colchicums than bees or flies, but they are nonetheless important pollinators for some plants. Except for a few exceptions (Masarinae, pollen wasps) adult wasps feed solely on nectar and do not collect pollen for their offspring, opting instead for other arthropods as a source of protein for developing larvae. Adult wasps, like bees, require carbohydrates to fuel their active lives and nectar fills this requirement. Wasps aren't as picky about the quality of nectar as bees are, and may be more willing to visit plants that produce nectar of a lower quality. I suspect that is the case here, considering the level of rainfall in this part of Scotland could have raised the level of the nectar to a level easily accessible to this unspecialized nectar-drinker, simultaneously diluting it.

Colchicum cilicicum with Apis mellifera

Colchicum cilicicum is native to Turkey, Syria, and Lebanon in forest margins, rocky slopes, and stream banks, often growing in limestone. It is probably one of the more commonly available species in the horticultural trade, which probably results from its ease of growth. I find this species to be the most reliable performer in my garden, and while it has smaller flowers than the others, it is my favorite. It makes up for the smaller size of its flowers by sending up many of them, and clumping readily. Add to this the long bloom period, about a month for this single clump, even though individual flowers only last a few days. The flowers are short, which means they are less easily knocked down.

Colchicum cilicicum with Halictus ligatus

Various bees and flies have visited the flowers of C. cilicicum, as well as carpenter ants (Camponotus sp.) seeking the sugary nectar. Pollination is always achieved by some means, most likely by insects, as apparent by the full seed pods it sends up every year.

I hope you will give this genus a try in your own garden, or find a way to see them in the wild. Corms are usually available for sale in the fall along with other spring blooming bulbs, corms, and tubers (See my Links for a few sources)

References

Dafni, Amots. "Autumnal and winter pollination adaptations under Mediterranean conditions." Bocconea. 5.171-181 (1996)
http://www.herbmedit.org/bocconea/5-171.pdf

Gómez-García, Daniel, José Azorín , Stella M. Giannoni and Carlos E. Borghi . "How does Merendera montana (L.) Lange (Liliaceae) benefit from being consumed by mole-voles?." Plant Ecology. 172.(2004): 173-181.
https://eurekamag.com/pdf.php?pdf=004187782

Kleizen, Ciara, Jeremy Midgley and Steve D. Johnson. "Pollination Systems of Colchicum (Colchicaceae) in Southern Africa: Evidence for Rodent Pollination." National Center for Biotechnology Information. Ann Bot. 2008 Nov; 102(5): 747–755
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2712380/

Piling, David, et al. "Colchicum." PBS Wiki. Pacific Bulb Society, 22 Jan 2016. Web.
http://pacificbulbsociety.org/pbswiki/index.php/Colchicum

Pisanty, Gideon and Yael Mandelik . "Effects of alien species on plant-pollinator interactions: how can native plants adapt to changing pollination regimes?." Evolution of Plant–Pollinator Relationships. Cambridge University Press. Chapter 15.(2012): 414-438.
https://www.researchgate.net/publication/266389244_Effects_of_alien_species_on_plant-pollinator_interactions_how_can_native_plants_adapt_to_changing_pollination_regimes

Rader et al.. "Non-bee insects are important contributors to global crop pollination." Proceedings of the National Academy of Sciences. 113.1 (2015):
https://www.researchgate.net/publication/285236115_Non-bee_insects_are_important_contributors_to_global_crop_pollination

Sütlüpinar, N. "The evaluation of Turkish autumn flowering Colchicum species." Journal of Faculty Pharmacy of Istanbul University 32 (1998): 1-8.

Young, Ian. "Bulb Log 36." SRGC Bulb Log Diary. Scottish Rock Garden Club, 8 Aug 2004.
http://www.srgc.org.uk/bulblog/log2004/300804/log.html

Young, Ian. "Bulb Log 37." SRGC Bulb Log Diary. Scottish Rock Garden Club, 13 Sep 2006.
http://www.srgc.org.uk/bulblog/log2006/130906/log.html

Autumn Bee Plants

noreply@blogger.com (Travis Owen) — Fri, 09 Sep 2016 13:23:00 +0000

A male long horned bee (Melissodes sp.) perched on Madia elegans with a face full of pollen.

Autumn is here, the days are warm, the nights are cool, and bees and other pollinators are still active. For me this is one of the best times of the year since we can finally get some relief from the intense heat of summer, and the leaves begin to change color on the trees. For bees, particularly honeybees, it is a crucial time to collect as many resources as possible so they can survive the winter. For solitary bees and other pollinators, most of which do not live in colonies or over winter as adults, nutrition is kept up in mating individuals and stores for brood are collected so the species may live on another year. For many of us gardeners, we love to see and support these creatures. This means reducing or eliminating the use of pesticides and providing them with good nutrition: flowers.

Apis mellifera queen mating yard flanked with Madia elegans

As a general rule, native flowering plants (although occasionally less showy) are far more useful to pollinators than exotic plants since the native pollinators have coevolved with them since time immemorial. Foreign plants, even those from a different region of the same state, generally do not attract the same diversity of pollinators, however, they may be highly attractive to a select few pollinators. I personally am not "strictly native" when it comes to gardening, so I grow many nonnative plants. I do appreciate the value and importance of natives, having seen firsthand the pollinators they attract that aren't attracted by exotic plants. This, in a sense, strikes a balance between gardening for conservation and gardening for aesthetics. Of course, for me, I am always gardening for pollinators.

Madia elegans

Madia elegans is one native wildflower that is just beginning to enter gardens. The flowers are open from dusk til dawn, and will remain open longer if grown with a Northern exposure or on cloudy days. They are highly attractive to honeybees and a variety of other bees and other pollinators, though for conservative gardeners they may appear a bit weedy and untidy.

Read more about Madia elegans and its various pollinators (Owen 2016)

Euthamia occidentalis with Halictus ligatus

Euthamia occidentalis is a native golgenrod, occasionally sold in nurseries, and is used for erosion control. It can be grown in damp places, something I have seen firsthand on the bank of the Rogue River. Like the true goldenrods (Solidago spp.) they are highly attractive to bees, as well as various wasps. I observed honeybees and halictid bees visiting the flowers. Sphecid wasps, Prionyx sp. I think, were also seen. Species of Prionyx paralyze grasshoppers as live food for their offspring in underground brood nests.

Cichorium intybus

Chicory is one of those nonnative plants that is found nearly everywhere humans are, a truly successful species. They are closely related to lettuce, and like lettuce have flowers that are open for a single day (often only in the morning) before withering. New flowers open daily, and attract honeybees and other bees, as well as other pollinators (like grass skippers, Hesperiinae).

Cichorium intybus (2015)

Being a composite, each stalk seen in this photo is an entire flower complete with staminate and pistillate parts, while the "petals" are actually sterile ray florets whose purpose is to attract and provide a landing pad to pollinators.

Read more about Cichorium intybus (Street et al. 2013)

Centaurea solstitialis and Apis mellifera

Related to chicory, in the sunflower family (Asteraceae), yellow starthistle is another nonnative plant which is adored by honeybees and beekeepers alike. Unlike chicory, which is unofficially considered to be a benign exotic by most, Centaurea solstitialis is on the noxious weed list in Oregon and at least ten other states and two Canadian provinces. Despite this, it is loved by beekeepers since it gives honeybees a lot of resources to make it through winter, since this region is generally considered to be in a dearth of nectar from mid-summer onward. I challenge this view, considering there are at least a few other plants, native and exotic, which are worked by bees and flower en masse in the latter half of the year (i.e. Madia elegans & Hypochaeris radicata, the latter another noxious nonnative adored by bees).

An apiary inundated with yellow starthistle

Yellow starthistle is native to southern Europe and western Eurasia, and in some parts of its native and introduced range is considered to be a supreme honey plant as the honey produced from it is one of the most sought and most popular varietal honeys according to some sources. The plants are self-incompatible, thus requiring insect pollinators. Honeybees account for most of the seed set, bumblebees following in importance. The apiary above was surrounded by a portion of uncut starthistle in Ashland, Oregon. The hills of Ashland are completely filled with this invader, showing the true destructive nature of noxious weeds, whose extreme competition outcompete native wildflowers and disrupt the natural balance. And in case you were wondering, I had to work all fifty+ hives in this apiary, and thanks to the thorny inflorescences, it was quite literally a pain in the ass (among other places).

Read more about Centaurea solstitialis (Zouhar 2002)

Cirsium vulgare

True thistles, such as bull thistle (Cirsium vulgare) are considered noxious weeds in Oregon and several other states. It is found in all fifty states of the US, yet native to Europe, Eurasia, and Africa. The plants are worked by honeybees and bumblebees, possibly others, though they often grow in very low density in this area so are not worked judiciously. They are known to grow in dense stands and I would presume that they would be quite covered in bees. The thorns are incredibly sharp and easily penetrate clothing and skin, which I can say is quite painful!

Lathyrus latifolius with a small leaf-cutter bee (Megachile sp.)

Another nonnative weed, quite widespread, the perennial sweet pea was once introduced to prevent erosion in ditches and banks. Fortunately, it is worked by a variety of bees including honeybees, bumblebees (Bombus spp.), large carpenter bees (Xylocopa spp.), and leaf-cutter bees (Megachile sp.). Peculiar, this nonnative pea is the only flower I have seen the leaf-cutters on. Except for the carpenter bees, all the bees I have seen appear to rob nectar by accessing it from above the nectaries rather than passing through the anthers, though these plants may very well be self-compatible like the garden pea since they seem to produce full pods most of the time (if not all the time).

Verbascum olympicum

Mullein (Verbascum spp.), occasionally "cowboy toilet paper" (I haven't, so don't ask), is an ubiquitous roadside weed, probably all over the country. Typically when one speaks of mullein one is speaking of Verbascum thapsus, or common mullein, which has a single flower spike and can reach seven or eight feet in height. A less common species, at least in my area, V. olympicum differs from V. thapsus most obviously by the multi branched inflorescence atop the plant, which attains a similar height. The individual flowers of V. olympicum appear to be slightly larger than those of V. thapsus, and the flowers of the latter seem to close or wither quicker than the former.

Verbascum olympicum and a hidden bumblebee

Both V. thapsus and V. olympicum, in my experience, are decent bee plants, though V. thapsus seems to only be worked in the morning when new flowers open, while V. olympicum seems to be worked all day. Another weedy mullein commonly encountered, V. blattaria, or moth mullein, is only occasionally worked by bumblebees but rarely honeybees, or at least I haven't seen it. All are grown in gardens, sometimes for their medicinal properties (the flowers of V. olympicum and V. thapsus are used for tea, and the plants are a source of water soluble saponins and mucilage).

Oenothera elata ssp. hookeri

The tall evening primrose, similar to some of the species native to Oregon, is one of many species grown in gardens. As the name implies, the plants can get really tall, up to four feet in the case of the plant photographed. The flowers open at dusk and remain open until morning when they wither. Though each flower is only open one night, new flowers open daily. They are primarily pollinated by large crepuscular sphinx moths (Sphingidae), some have visited this plant on more than a few occasions though I was unable to attain a photo. Honeybees adore it, as you can probably tell, and can be seen visiting the flowers as they open on warm nights, or in the morning before the flowers wither. There are hybrids of some of the smaller oenotheras available (i.e. Oenothera kunthiana), though they do not appear to be as attractive to pollinators in my garden.

Asclepias fascicularis and Apis mellifera

Many milkweeds are useful bee plants, though the flowers of some species can pose a threat to bees and other pollinators (more on that later). Milkweeds are important honey plants in much of the US where they are common. They are also paramount to the survival of many species of butterflies, including the monarch (Danaus plexippus) and other milkweed butterflies (Danainae) since the caterpillars of these butterflies are completely dependent on them for sustenance. Certain species of moths and a few true bugs are also dependent on milkweeds for survival.

Read more about milkweeds as honey plants (Krochmal 2016)

Apis mellifera with pollinia on her hind foot

Milkweeds are peculiar plants for a variety of reasons. For an amateur botanist, I find their breeding system intriguing. The most notable feature of this, I think, is that the pollen is in the form of pollinia, as is found in orchids, where the pollen from each anther is composed into a single sticky mass. This mass adheres itself to the foot, or anywhere on the pollinator before being hooked onto the "claws" surrounding the pistil. Some unfortunate pollinators whose feet have pollinia attached become stuck themselves on these hooked appendages (called horns) and not having the strength to escape, die of dehydration or something similar. Butterflies, often being larger and more powerful fliers, can probably escape the horns quite easily, but bees, beetles, and wasps have been found hanging, dead, on the flowers of some species.

Read more about the Fatal Attraction of Milkweeds (Chen 2006)

Croton setigerus

In dry sunny fields, an inconspicuous plant in the Euphorbiaceae forms tight grey-green cushions of hairy foliage and tiny inconspicuous flowers. This is Croton setigerus, or turkey mullein, found in much of the Western US in dry hot sites. It is occasionally worked by honeybees, and claimed to produce a "thick amber honey."

Read more about Croton setigerus (Owen 2016)

Trichostemma lanceolatum

Trichostema is a genus in the Lamiaceae, or mint family. A few of which are notoriously bee pollinated, such as the perennial California native T. lanatum, woolly blue curls, which is widely proclaimed to be a good honey plant. In my region, the annual T, lanceolatum is a somewhat locally common plant that grows in poor soil with low grasses and other small forbs. In my experience, it is more frequently visited by skippers (Ochlodes sylvanoides) who appear to be a good fit for the plants highly elongated reproductive structures.

Ochlodes sylvanoides on Trichostema lanceolatum

Pollinator visitation, it seems, is a highly regionally specific variable. This has been seen in many other plants, and can readily be studied by any gardener who grows nonnative plants. In the case of Trichostema, where I have witnessed butterflies, many other people have reported seeing various bees visit the flowers of T. lanceolatum and there is similar literature on bee pollination of T. lanatum. Aside from bees, hummingbirds have been observed on some species, probably another regionally variable observation. To me, this speaks of the adaptability of some species to overcome the boundaries of others, in this case perhaps an inadequate supply of bees (not specifically due to bee losses, but maybe the normal ebb and flow of native bee activity in this area, correlating with the diversity, or lack thereof, of flowering plants at any given time).

Read more about Trichostema lanceolatum (Owen 2016)

Nepeta cataria and Bombus vosnesenskii

In the garden, mints and their relatives are a sure thing when it comes to bee plants. I do grow a variety of mints, all in one corner of the yard since they can be so invasive, and the flowers do attract a variety of bees. I prefer the more well behaved mint relatives, such as Agastache and Nepeta. Common catnip is a short lived perennial that has survived with little care at the border of my garden for four years now. It was started by seed scattered onto disturbed soil and watered for the first year. Now it is four feet tall and covered in small flowers. Many species of native bee are attracted to the flowers, such as Lasioglossum and Bombus vosnesenski. Honeybees also adore the flowers, there may be upwards of fifty bees on this single plant at any one time.

Thymus serpyllum 'Elfin' with Apis mellifera

Another "mint," thyme is always attractive to honeybees. The particular selection T. serpyllum 'Elfin' appears to be later blooming than the others in my garden, and is an excellent plant in my small rock garden. Thyme is the source of the thymol, an antimicrobial compound used for the control of varroa mites (Varroa destructor) among other things. There is a folk tale that where thyme is grown for seed at the field scale the honeybees nearby are free of mites. I cannot speak to the legitimacy of this claim, but the notion that very small quantities of thymol or other mite-repellent compounds are found in the nectar is not without merit.

Aerial yellowjacket (Dolichovespula arenaria), bald-faced hornet (D. maculata)

While talking about autumn bee plants, it is worth noting some of the bees cousins that are also highly active at this time. Aerial yellowjackets and bald faced hornets (both in the genus Dolichovespula) are predatory social wasps which build enclosed paper nests in trees or sometimes under overhangs of human made structures. At best, they are predators of garden pests such as tomato hornworms (Manduca quinquemaculata) and others, but at worst they are bee predators who can destroy weak honeybee hives. For strong hives they are merely a nuisance.

Practices to help your hives protect themselves include using the smallest opening of an entrance reducer, maybe a one inch entrance hole, and feeding your bees (inside the hive is best, i.e. hive top feeders or some variant) so they build in number and strength to ward off any potential intruders. Yellowjacket traps can be useful where they are very populous, but I would suggest using a hot dog or lunch meat as bait rather than the commercially available bait. I don't usually condone the killing of hymenopterans, even wasps, but to be sure with simple traps (a chemical free approach) their populations will not be devastated and they will probably bounce back the following year.

When your cousin shows up at the party uninvited...

...and disembowels your family.

It happens.

Colchicum 'Lilac Wonder' with a carpenter ant (Camponotus sp.)

To avoid ending on a gruesome note, here is one more good group of bee plants: Colchicum. Sometimes referred to as the autumn crocus, this name is a poor representation of the genus since there are autumn, winter, and spring blooming species in both Colchicum and Crocus. Colchicum, unlike Crocus, grows from a toxic structure that is like something between a bulb and a corm, quite unique, although typically considered to be a corm. Crocus on the other hand grows strictly from a corm and is not toxic, in fact it is practically edible and it is eaten by rodents, deer, turkeys, and slugs. Colchicum is virtually pest free, due to the toxins within the plant. Most of the commercially available colchicums are the autumn blooming types. These tend to flower in the fall, and send up leaves and seed pods in spring. I've grown colchicums for three years, and honeybees seem to adore them.

Colchicum cilicicum

I grow four varieties including three fall blooming and one spring blooming although the spring blooming type hasn't grown to flowering size yet. The first variety I attained was the hybrid C. 'Lilac Wonder,' a commonly available and very popular large flowered hybrid supposedly between C. giganteum and C. bornmuelleri (both sometimes considered synonyms of C. speciosum), bred sometime in the early 1900s. I also grow C. speciosum, which has slightly smaller flowers and is shorter, but equally if not more beautiful.

Colchicum cilicicum

Of the few colchicums that I grow, C. cilicicum is by far the most floriferous and longest blooming, thus the best for bees. The individual flowers don't last very long, but C. cilicicum sends up new flowers in succession for a few weeks. Despite there being so few of these flowers, there is a continuous stream of honeybees visiting them for both pollen and nectar. I imagine a field full of Colchicum would be quite active with bees of various types. I would do just that, but the cost of these large bulb/corms is somewhat prohibitive for me. I am attempting a few species from seed, and have a hopeful view of the future.

Oenothera elata ssp. hookeri with six honeybees

There are many other fall blooming plants that may be excellent bee plants. Some recommended by others include Aster spp., Caryopteris spp., Clematis ternifolia, Epilobium spp., Eutrochium spp., Hylotelephium spectabile, Lythrum salicaria, Perovskia atriplicifolia, Pycnanthemum flexuosum, Rosmarinus spp., Tetradium daniellii, Vernonia altissima, Vitex agnus-caste, and no doubt many others. I'd love to hear about more bee plants, put them in the comments (with your location for context)!

The Amateur Anthecologist

Andrena: A Closer Look

Bumble Bees of Vesper Meadow

Vesper Meadow Restoration Preserve

Bombus vosnesenskii on Solidago at Vesper Meadow2021 Bumble Blitz at Vesper Meadow

Bombus flavifrons on Agastache urticifolia

Bumble Bees 101

Bombus californicus on Wyethia angustifoliaBumble Bee Life Cycle

Bombus bifarius s.l. (B. vancouverensis nearcticus)Vesper Meadow Bumble Bees

Species Accounts

White-Shouldered Bumble Bee - Bombus appositus

California Bumble Bee - Bombus californicus

Yellow-Fronted Bumble Bee - Bombus flavifrons

Fuzzy-Horned Bumble Bee - Bombus mixtus

Black-Tailed Bumble Bee - Bombus melanopygus (syn. edwardsii)

Nevada Bumble Bee - Bombus nevadensis

Two-Formed Bumble Bee - Bombus bifarius s.l. (syn. vancouverensis nearcticus)

Yellow-Faced Bumble Bee - Bombus vosnesenskii

(including descriptions for vandykei and caliginosus)

Learn more about Bumble Bees:

Table Rock Revisited

Floral Visitors MMXXI

I Miss Summer: A Soliloquy

Summer Bees, Wasps, and Other Pollinators

Pollinators and Wildflowers of Mount McLoughlin

More Bees from Spring '18

Bees in Spring 2018

Arctostaphylos viscida

References:

Parasitoid Wasps of Southern Oregon 2017

Ichneumonoidea

Ichneumonidae

Ichneumoninae

Banchinae

Cryptinae

Braconidae

Evanioidea

Gasteruptiidae

Cynipoidea

Cynipidae

Chalcidoidea

Torymidae

Pteromalidae

Diaprioidea

Diapriidae

Conclusion

References

Useful Websites

Aculeate wasps of Southern Oregon pt 2

Sphecidae

Pompilidae

Mutillidae

Sapygidae

Chrysididae

Bethylidae

Tiphiidae

Aculeate Wasps of Southern Oregon pt. 1

Eusocial Vespidae

Solitary Vespidae

Crabronidae

Apidae in Southern Oregon

Daucus carota

Thoughts on Rabbitbrush and Other Things

Trichostema lanceolatum

Mount McLoughlin

Lower Table Rock

Moths and Butterflies of Southwestern Oregon (2016)

Superfamily Alucitoidea

Family Alucitidae

Superfamily Lasiocampoidea

Family Lasiocampidae

Subfamily Macromphaliinae

Superfamily Noctuoidea

Family Erebidae

Subfamily Arctiinae

Family Noctuidae

Subfamily Amphipyrinae

Subfamily Noctuinae

Subfamily Plusiinae

Superfamily Pterophoroidea

Bombus vosnesenskii on Solidago at Vesper Meadow
2021 Bumble Blitz at Vesper Meadow

Bombus californicus on Wyethia angustifolia
Bumble Bee Life Cycle

Bombus bifarius s.l. (B. vancouverensis nearcticus)
Vesper Meadow Bumble Bees