Halacaridae is a family of meiobenthic
mites found in marine, brackish, and freshwater habitats around the world. It includes more than 1100 described species belonging to 64 genera [1][2] It is the largest marine radiation of arachnids.[3]
Description and life cycle
Halacarids have four pairs of legs (as adults and nymphs; see below), of which the first two pairs point forwards and the last two pairs point backwards. This is a
synapomorphy of the group. Another synapomorphy are four plates on the dorsal surface of the body, except for several genera with reduction of certain plates.[4]
The life cycle of halacarids consists of egg,
larva, 1-3
nymphal stages (protonymph->deutonymph->tritonymph) and adult. Additionally, between each of the free-living stages (i.e. except for the egg) is a quiescent pupal stage.[5]
Eggs are usually deposited by an adult female in a substrate, with the help of an
ovipositor.
Larvae have three pairs of legs, with each leg five-segmented, and lack a genital plate.
Protonymphs have four pairs of legs (as do all following stages), of which the fourth pair is five-segmented, and they usually have a distinct genital plate.
Deutonymphs and tritonymphs have each leg six-segmented.
Adults are often similar to the last nymphal stage, but they have an ovipositor (if female) or spermatopositor (if male).
Like mites in general, halacarids have a pair of
palps. The palps usually have four segments each, but they are three-segmented in Simognathus and just two-segmented in Acaromantis.[3]
Halacarids of subfamily Copidognathinae have just a single nymphal stage. Additionally, the number of
genital papillae is reduced to a single pair.[3]
The subfamily Rhombognathinae, which is
algivorous, can be recognised by the dark green or almost black pigment inside their digestive system. This pigment is partially digested
chlorophyll from algae.[3]
Ecology
Halacaridae occur in various habitats including sandy beaches, tidal sediment, interstitial spaces,
hydrothermal vents,
mangroves,
salt marshes and on larger animals.[2][6] They spend their entire lives on a substrate such as attached
algae or sand.[3]
Most species and genera are
predators, though Rhombognathinae are instead algivores.[3] In freshwater halacarids, some species are restricted to
crayfish gill chambers, implying a
parasitic lifestyle, while Lobohalacarus weberi is a
scavenger that feeds on dead
nematodes and
oligochaetes but not on live ones.[7]
Phylogeny
Recent analyses place Halacaridae as the
sister group to
Parasitengona.[8][9] Within the group, algivorous Rhombognathinae consists of two lineages (Rhombognathus+Isobactrus and Rhombognathides+Metarhombognathus), meaning the habit of algivory has evolved two independent times.[3]