phoresy

Who likes shrews? These mites do!

My friend Allan Lindoe, fossil preparator extraordinaire, lives on an acreage near Athabasca and makes the journey south to Edmonton about once a week to carefully remove rocky matrix from around the skeletons of long-dead fishes, mosasaurs, and dinosaurs. Two cats share his home and frequently bring him presents of wild game. A few weeks ago I washed a mixed bag (literally) of a dozen shrews he had accumulated over the summer and fall of 2015. Chewed-on shrews are not easy to identify unless you know a lot about insectivore teeth, but based on tail length and known distributions of shrew species in Alberta, they were one or more of the masked shrew (Sorex cinereus), Arctic shrew (S. arcticus), pygmy shrew (S. hoyi) and/or dusky shrew (S. monticolus)*. Some of the shrews were rather decomposed, so I wasn’t expecting much from the washings, but I was pleasantly surprised: 6 species of mites! Members of both major lineages were present. From the Acariformes were Prostigmata (Myobiidae, Pygmephoridae and Trombiculidae) and Sarcoptiformes (Glycyphagidae). From the Parasitiformes there were larval hard ticks (Ixodida: Ixodidae) and what look like Melicharidae (Mesostigmata).  Myobiids, trombiculids, and ixodids are parasitic, and the others are all likely just phoretic. Who knew the zoo on shrews? Now you do.

Protomyobia female ex shrew

Protomyobia nr. claparedei female (note egg).

 

Protomyobia ex shrew

Protomyobia nr. claparedei male (note well-sclerotized aedeagus).

 

Protomyobia lv I think A

Larval Protomyobia nr. claparedei (note styletiform chelicerae).

 

pygmephorid ex shrew

Ventral view of one of the many pygmephorids from the shrew washings.

 

pygmephorid ex shrew legs I

Pygmephorid showing modified legs I.

 

chigger ex shrew A

Dorsal view of a shrew chigger (Trombiculidae).

 

chigger ex shrew B

Prodorsal shield of a shrew chigger, showing the posterior pair of trichobothria and single anterior median seta (just the base can be seen here) indicative of the family Trombiculidae, as opposed to members of the Leeuwenhoekiidae, which have two anterior median setae.

 

prob Oryctoxenus A

An Oryctoxenus sp. deutonymph (Glycyphagidae). Anterior is pointing up, and yes, it doesn’t have mouthparts.

 

prob Oryctoxenus B

Posterior hair-clasping structures of an Oryctoxenus deutonymph.

 

ixodid larva ex shrew A

A larval hard tick (Ixodidae).

 

ixodid larva ex shrew B

Retrorse spines on the tick’s hypostome help keep it attached to the host.

 

maybe Proctolaeleps A

Maybe a female Proctolaelaps sp.(Melicharidae). Not in great shape.

 

maybe Proctolaeleps B

The ‘procto’ part of Proctolaelaps refers to the large anal opening, or so the etymological legend goes.

*Smith, H.C. 1993. Alberta Mammals: an Atlas and Guide. The Provincial Museum of Alberta, Edmonton, Alberta.

 

 

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Is That a Wombat on Your Belly, Or Are You Just Happy to See Me?

I attended a Ph.D. defense a few weeks ago on the effects of salmon lice (which are copepods, not insects) on their juvenile hosts. The student showed some gory photos and pointed out that for such a little fish, carrying a louse was like a human lugging around a raccoon on his back. Some mites can be just as burdensome, such as this Macrocheles muscaedomesticae (Scopoli) attached to the abdomen of an unfortunate Drosophila hydei Sturtevant.

Scanning electron micrograph of a Drosophila hydei carrying a female Macrocheles muscaedomesticae (image by HP)

Scanning electron micrograph of a Drosophila hydei carrying a female Macrocheles muscaedomesticae (image by HP)

Females of many species of Macrocheles (Mesostigmata: Macrochelidae) hitch rides on winged insects to move from a place to place, a phenomenon called phoresy. Strictly phoretic organisms do not feed on the host while attached. A great many mite taxa fall into this ‘purely phoretic’ category. Others may facultatively snack on the host while in transit. My colleague Lien Luong has investigated one such mite species, Macrocheles subbadius (Berlese), and its cactus-associated host Drosophila nigrospiracula Patterson & Wheeler. When Lien moved to the University of Alberta it proved difficult to replicate the system, in part because cacti are not common in this part of Alberta.  Compost bins are abundant, however, and Lien and her students are investigating the ecological relationship between two compost-associated species, Macrocheles muscaedomesticae and Drosophila hydei. Does M. muscaedomesticae feed on its host while attached, or is it just holding on? One way to test this is to determine whether the mite’s mouthparts pierce the fly’s integument. In this N = 1 sample, the mite just seems to be holding on firmly, probably uncomfortably so from the fly’s point of view.

mite biting medium close

Piercing or just pinching?

 

mite biting close

Looks like pinching, probably painfully.

But by definition, facultative parasites aren’t always parasitic. More mites must be examined, and other lines of evidence followed, such as presence of melanized wounds on hosts after the mites have dropped off, or presence of Drosophila DNA in the guts of the Macrocheles.

 

On the backs of wasps

In March, I was given two specimens of solitary wasps that were covered with mites. The first was one of several Crossocerus  (Crabronidae) that had overwintered in holes in a wooden chair left outside on the campus of the University of Alberta. I had expected the mites to be phoretic deutonymphal astigmatans, but they weren’t, they were adult female scutacarids (Prostigmata: Scutacaridae). Many scutacarid species have phoretic and non-phoretic morphs. The big anterior tarsal claws you can see (blurrily) on the photo below are typical for phoretomorphs. What wasn’t typical was a pair of strange internal structures that became apparent in well-cleared specimens.

scutacarid from Crossocerus April 2015 E

At first I thought the pair of round things near the female’s genital area were sperm-storage chambers. But when I Googled ‘Crossocerus’ and ‘Scutacaridae’, I found a paper that showed I was only half right – they were sporothecae*, not spermathecae!

scutacarid from Crossocerus April 2015 C

Two big spores tucked into the genital atrium of this female Imparipes.

scutacarid from Crossocerus April 2015 A

Ebermann & Hall (2004) described a new species of scutacarid, Imparipes haeseleri, from several species of wood-associated Hymenoptera. In the genital atrium of these mites, they observed two large round fungal spores, one on each side, looking remarkably similar to the ones in the mites from the rotting chair. I asked Evert Lindquist, an expert on the Heterostigmata (the larger group to which Scutacaridae belongs) if these mites were Imparipes. Yup, they were. Were they I. haeseleri? There is a closely related species known from North America, I. vulgaris, but several setal characters matched haeseleri rather than vulgaris so I decided to go with Imparipes cf. haeseleri.

Why are the female mites carrying spores? No doubt they and their offspring feed on the wood-digesting mycelium produced from the germinated spores. The mites that hop on wasps as they depart from their overwintering chambers take with them the starter culture for their future meals. Dr. Lindquist noted that the spores these mites were carrying looked very similar to the Nigrospora spores known to be carried by a different species of heterostigmatan, Siteroptes reniformis Krantz. In his 1984 paper, Lindquist notes that S. reniformis “not only serve to transport and place spores of Nigrospora in an environment favorable for germination and growth, they also stimulate mycelial growth, apparently by secreting a chemical substance when feeding on the fungus.”

The second wasp was collected from an overwintered artificial nesting block that was supposed to house solitary bees. It was an Ancistrocerus sp. (Vespidae: Eumeninae). Knowing this, it was easy to guess who the mites were, and slide-mounting confirmed it: deutonymphs of a Kennethiella sp. (Astigmata: Winterschmidtiidae).

Winterschmidtiid_001

Like almost all phoretic deutonymphs of Astigmata, these Kennethiella have a terminal sucker plate to adhere to hosts. Unusually, they also have anterior ocelli. Why ocelli are present in only a small number of Astigmata is unclear (at least, it’s unclear to me).

Winterschmidtiid_003

Sucker plate.

Winterschmidtiid ocelli

Pair of ocelli.

The reason I expected the mites to be Kennethiella is because the relationship between them and their host wasps is famous among acarologists. Cowan (1984) unraveled the interactions for one mite-wasp duo.  To quote the abstract: “The mite Kennethiella trisetosa is phoretic on adults of the wasp Ancistrocerus antilope and develops in the nest with immature wasps. Female mites and a large type of male develop oviparously, whereas a small male develops oviparously. Small males kill each other, but are ignored by large males. By mating with females before small males are mature, large males may monopolize fertilization. Larvae of female wasps usually destroy mites within their cells but, as adults, are reinfested when mated by mite-bearing males. Each time a male wasp mates, about half of its mites transfer to the female.”

It’s worth reading the original to appreciate the full intricacies of these intertwined life-cycles.

Ancistrocerus showing Kennethiella mites 16 Aug 2009

A home-grown Ancistrocerus with a load of Kennethiella, from my back yard in Edmonton a few years ago.

*according to Evert Lindquist, they aren’t sporothecae (which are spore-storage sacs) but simply the spores themselves, tucked into corners of the genital atrium. Thanks, Evert!