Corresponding author: Ashley P. G. Dowling (
Academic editor: V. Pesic
Thirteen new species of North American
O’Neill JC, Fisher JR, Nelson WA, Skvarla MJ, Fisher DM, Dowling APG (2016) Systematics of testudacarine torrent mites (Acari, Hydrachnidia, Torrenticolidae) with descriptions of 13 new species from North America. ZooKeys 582: 13–110. doi:
Despite their abundance, few testudacarines are described worldwide and in North America the most recent description is over fifty years old. Limited morphological and distributional data have been presented, and no genetic data has ever been published on
Thirteen descriptions and four redescriptions of North American
There are currently nine testudacarines described worldwide:
Only two authors,
Mites were collected and preserved using protocols detailed in
Terminology used in this study is detailed in Figs
Testudacarine male dorsum (generalized): (
Testudacarine male dorsum (SEM): anterio-medial platelet (amp); anterio-lateral platelet (alp); dorsal plate (dp); dorso-glandularia (dg); post-ocularial setae (pos); dorsal membrane (dm); latero-glandularia (lg). Scale: 100 µm. Photo Michelle Hoppner and Ian Smith (used with permission).
Testudacarine male venter (generalized):
Testudacarine male venter (SEM): coxo-glandularia (cg); latero-glandularia (lg); ventro-glandularia (vg); coxae (c); coxae-II+III midline (ml); genital field (gf); acetabula (a); line of secondary sclerotization (ss); excretory pore (ep). Scale: 100 µm. Photo Michelle Hoppner and Ian Smith (used with permission).
Testudacarine gnathosoma (generalized): chelicerae (c); posterio-dorsal apodeme (pda); posterio-ventral apodeme (pva); subcapitulum (sc); fang (f); rostrum (r).
Images were produced and measurements taken following the protocol detailed in
All holotypes, allotypes, and some paratypes have been deposited in the Canadian National Collection of Insects, Arachnids, and Nematodes
Material from the
The “barcoding” region of COI was used as an independent test of morphological species hypotheses. COI was used to determine if any morphological characters, conservative or ambiguous, indicated species boundaries by sorting into distinct genetic lineages. COI was also used in the same way to test distributional hypotheses. Taxon sampling included roughly 300 specimens spanning “morphotypes” from across North America. Unfortunately, ethanol collections were limited from Mexico, northern Canada, and the eastern United States and therefore do not fully represent the ranges of species from these regions. Later, twenty specimens were included for phylogenetic analysis of 28S (D1-3) to investigate interspecific relationships. Genbank accession numbers of specimens for which sequences were obtained and used in this study are located in Table
Genbank accession numbers and GenSeq nomenclature for each specimen sequenced for this study.
Species | Genbank Accession # | Specimen Catalog # | ||
---|---|---|---|---|
COI | 28S | |||
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ACUA135545 (non-type voucher) | genseq-4 COI | |
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ACUA135544 (non-type voucher) | genseq-4 COI, 28S |
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ACUA135543 (paratype) | genseq-2 COI | |
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ACUA146756 (paratype) | genseq-2 COI | |
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ACUA138471 (paratype) | genseq-2 COI | |
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ACUA141898 (holotype) | genseq-1 COI | |
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ACUA138472 (paratype) | genseq-2 COI | |
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ACUA138473 (paratype) | genseq-2 COI | |
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ACUA138469 (paratype) | genseq-2 COI | |
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ACUA144021 (paratype) | genseq-2 COI | |
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ACUA138476 (non-type voucher) | genseq-4 COI | |
|
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ACUA141897 (holotype) | genseq-1 COI | |
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ACUA138476 (non-type voucher) | genseq-4 COI | |
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ACUA138478 (non-type voucher) | genseq-4 COI | |
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ACUA141903 (non-type voucher) | genseq-4 COI | |
|
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ACUA138479 (non-type voucher) | genseq-4 COI | |
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ACUA141904 (non-type voucher) | genseq-4 COI | |
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ACUA138480 (non-type voucher) | genseq-4 COI | |
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ACUA138481 (non-type voucher) | genseq-4 COI | |
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ACUA138482 (non-type voucher) | genseq-4 COI | |
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ACUA141901 (non-type voucher) | genseq-4 COI | |
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ACUA141902 (non-type voucher) | genseq-4 COI | |
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ACUA141900 (non-type voucher) | genseq-4 COI | |
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ACUA138484 (non-type voucher) | genseq-4 COI | |
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|
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ACUA138487 (non-type voucher) | genseq-4 COI, 28S |
|
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ACUA141899 (non-type voucher) | genseq-4 COI | |
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ACUA141905 (non-type voucher) | genseq-4 COI | |
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ACUA138486 (non-type voucher) | genseq-4 COI | |
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ACUA138488 (non-type voucher) | genseq-4 COI | |
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ACUA138494 (holotype) | genseq-1 COI | |
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ACUA138495 (holotype) | genseq-1 COI | |
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ACUA138489 (non-type voucher) | genseq-4 COI | |
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ACUA141906 (non-type voucher) | genseq-4 COI | |
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ACUA138490 (non-type voucher) | genseq-4 COI | |
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ACUA138491 (non-type voucher) | genseq-4 COI | |
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ACUA138492 (non-type voucher) | genseq-4 COI | |
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ACUA138493 (non-type voucher) | genseq-4 COI | |
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ACUA143634 (paratype) | genseq-2 COI | |
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ACUA141892 (paratype) | genseq-2 COI | |
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ACUA141893 (paratype) | genseq-2 COI | |
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ACUA141894 (paratype) | genseq-2 COI, 28S |
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ACUA142194 (non-type voucher) | genseq-4 COI, 28S |
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ACUA141896 (paratype) | genseq-2 COI | |
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ACUA143618 (paratype) | genseq-2 COI | |
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ACUA143629 (paratype) | genseq-2 COI | |
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ACUA143633 (non-type voucher) | genseq-4 COI | |
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ACUA141895 (paratype) | genseq-2 COI | |
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ACUA143619 (paratype) | genseq-2 COI | |
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ACUA143623 (paratype) | genseq-2 COI | |
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ACUA141885 (holotype) | genseq-1 COI | |
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ACUA141886 (non-type voucher) | genseq-4 COI | |
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ACUA141887 (non-type voucher) | genseq-4 COI, 28S |
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ACUA142195 (non-type voucher) | genseq-4 COI | |
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ACUA141888 (paratype) | genseq-2 COI | |
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ACUA141889 (paratype) | genseq-2 COI, 28S |
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ACUA142196 (paratype) | genseq-2 COI | |
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ACUA142197 (paratype) | genseq-2 COI | |
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ACUA141890 (non-type voucher) | genseq-4 COI | |
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ACUA142198 (non-type voucher) | genseq-4 COI | |
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ACUA142199 (paratype) | genseq-2 COI | |
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ACUA142200 (non-type voucher) | genseq-4 COI, 28S |
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ACUA141891 (non-type voucher) | genseq-4 COI | |
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ACUA143643 (non-type voucher) | genseq-4 COI | |
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ACUA143644 (non-type voucher) | genseq-4 COI | |
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ACUA143645 (holotype) | genseq-1 COI | |
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ACUA143646 (non-type voucher) | genseq-4 COI | |
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ACUA143647 (non-type voucher) | genseq-4 COI | |
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ACUA143648 (paratype) | genseq-2 COI, 28S |
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ACUA143649 (paratype) | genseq-2 COI | |
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ACUA143652 (holotype) | genseq-1 COI, 28S |
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ACUA143654 (non-type voucher) | genseq-4 COI, 28S |
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ACUA143655 (non-type voucher) | genseq-4 COI | |
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ACUA143657 (non-type voucher) | genseq-4 COI | |
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ACUA143658 (non-type voucher) | genseq-4 COI | |
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ACUA143659 (non-type voucher) | genseq-4 COI | |
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ACUA143661 (non-type voucher) | genseq-4 COI | |
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ACUA143664 (non-type voucher) | genseq-4 COI | |
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ACUA143665 (non-type voucher) | genseq-4 COI, 28S |
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ACUA143666 (paratype) | genseq-2 COI | |
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ACUA143667 (non-type voucher) | genseq-4 COI, 28S |
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ACUA143669 (non-type voucher) | genseq-4 COI | |
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ACUA143671 (non-type voucher) | genseq-4 COI | |
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ACUA146717 (non-type voucher) | genseq-4 COI | |
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ACUA146718 (non-type voucher) | genseq-4 COI | |
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ACUA146719 (non-type voucher) | genseq-4 COI | |
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ACUA146720 (non-type voucher) | genseq-4 COI | |
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ACUA146721 (non-type voucher) | genseq-4 COI | |
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ACUA146722 (non-type voucher) | genseq-4 COI | |
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ACUA146723 (non-type voucher) | genseq-4 COI | |
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ACUA146727 (paratype) | genseq-2 COI | |
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ACUA146724 (paratype) | genseq-2 COI, 28S |
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ACUA146725 (holotype) | genseq-1 COI | |
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ACUA146726 (paratype) | genseq-2 COI | |
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ACUA146728 (holotype) | genseq-1 COI | |
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ACUA146729 (non-type voucher) | genseq-4 COI | |
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ACUA146732 (paratype) | genseq-2 COI | |
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ACUA146733 (non-type voucher) | genseq-4 COI | |
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ACUA146734 (non-type voucher) | genseq-4 COI | |
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ACUA146735 (non-type voucher) | genseq-4 COI | |
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ACUA146736 (paratype) | genseq-2 COI | |
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ACUA146738 (paratype) | genseq-2 COI | |
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ACUA146737 (paratype) | genseq-2 COI | |
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ACUA146739 (non-type voucher) | genseq-4 COI | |
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ACUA146740 (paratype) | genseq-2 COI | |
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ACUA146742 (paratype) | genseq-2 COI | |
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ACUA146743 (paratype) | genseq-2 COI, 28S |
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ACUA146744 (paratype) | genseq-2 COI | |
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ACUA146745 (paratype) | genseq-2 COI | |
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ACUA146746 (paratype) | genseq-2 COI | |
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ACUA146747 (paratype) | genseq-2 COI | |
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ACUA146748 (paratype) | genseq-2 COI | |
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ACUA146749 (non-type voucher) | genseq-4 COI | |
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ACUA146750 (paratype) | genseq-2 COI | |
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ACUA146751 (paratype) | genseq-2 COI | |
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ACUA146752 (holotype) | genseq-1 COI | |
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ACUA146753 (paratype) | genseq-2 COI | |
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ACUA146754 (non-type voucher) | genseq-4 COI | |
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ACUA146755 (paratype) | genseq-2 COI | |
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ACUA146756 (paratype) | genseq-2 COI | |
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ACUA146757 (paratype) | genseq-2 COI | |
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ACUA146758 (non-type voucher) | genseq-4 COI | |
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ACUA146759 (paratype) | genseq-2 COI | |
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ACUA146760 (non-type voucher) | genseq-4 COI | |
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ACUA146762 (holotype) | genseq-1 COI | |
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ACUA146763 (paratype) | genseq-2 COI, 28S |
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ACUA146764 (paratype) | genseq-2 COI | |
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ACUA146768 (non-type voucher) | genseq-4 COI | |
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ACUA146769 (holotype) | genseq-1 COI, 28S |
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ACUA146770 (paratype) | genseq-2 COI | |
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ACUA146772 (paratype) | genseq-2 COI | |
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ACUA146774 (paratype) | genseq-2 COI | |
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ACUA146775 (paratype) | genseq-2 COI | |
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ACUA146776 (paratype) | genseq-2 COI | |
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ACUA146777 (paratype) | genseq-2 COI | |
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ACUA146778 (non-type voucher) | genseq-4 COI | |
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ACUA146779 (non-type voucher) | genseq-4 COI | |
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ACUA146780 (non-type voucher) | genseq-4 COI | |
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ACUA146781 (non-type voucher) | genseq-4 COI, 28S |
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ACUA146778 (non-type voucher) | genseq-4 COI | |
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ACUA146782 (paratype) | genseq-2 COI, 28S |
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ACUA146783 (paratype) | genseq-2 COI | |
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ACUA146784 (paratype) | genseq-2 COI, 28S |
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ACUA146785 (non-type voucher) | genseq-4 COI | |
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ACUA146786 (non-type voucher) | genseq-4 COI | |
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No voucher | No classification |
Genomic DNA extraction was completed with Qiagen DNeasy Tissue Kits (Qiagen Inc.,Valencia, California). Amplifications of the target region of COI were performed with LCO1490 and HCO2198 (
Phylogenetic analysis of COI and 28S resulted in five well-supported (posterior probability greater than 95%) clades; however, analyses did not produce resolution at the base of
Three morphotypes (
In summary, we find strong support through a combination of morphology, biogeography, and phylogenetic analysis of COI and 28S for 17 species sorted into four robustly supported species complexes. The following species complexes are proposed to better organize the subfamily:
1 | Pedipalp four-segmented, anterior tips of coxae-I with projections |
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– | Pedipalp five-segmented, anterior tips of coxae-I without projections |
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2 | Body elongate to rectangular |
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– | Body oval |
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3 | Body large (>700 µm female and >650 male dorsal length), dull coloration common; within and west of the Rocky Mountains |
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– | Body small (<700 µm female and <650 male dorsal length), bright coloration (orange, red, violet, blue) common; present throughout North America | ( |
4 | Anterio-medial platelet wide (>140 µm) and more than or nearly twice as wide as long |
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– | Anterio-medial platelet unmodified (<140 µm) and far less than twice as wide as long |
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5 | Anterio-medial and anterio-lateral platelets with consistent coloration (either colored or colorless across) |
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– | Anterio-lateral platelets with coloration and anterio-medial platelet colorless |
|
See
For larval diagnosis see
Testudacarines have been reported on many occasions outside of their original descriptions. Furthermore, the Canadian National Collection in Ottawa, Canada includes thousands of testudacarines collected from across most of North America (
The three pairs of acetabula, coxae-IV condyles, and “generalized” pedipalps are plesiomorhphic states that clearly show testudacarines as retaining ancestral torrenticolid characteristics (
Testudacarine sexual dimorphism: female dorsal shield (
Known from only two counties (Los Angeles and Monterey) in California.
Lectotype (1♀), and paralectotype (1♂) deposited at
Gnathosoma (Fig.
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [459–505 total; trochanter 54–62; basifemur 81–91; telofemur 63–68; genu 81–91; tibia 86–100; tarsus 84–95]. Leg-II [516–554 total; trochanter 62–65; basifemur 85–100; telofemur 63–71; genu 84–96; tibia 100–114; tarsus 106–115]. Leg-III [593–644 total; trochanter 63–69; basifemur 97–105; telofemur 70–78; genu 104–118; tibia 125–143; tarsus 130–142]. Leg-IV [779–862 total; trochanter 84–96; basifemur 118–127; telofemur 115–129; genu 141–166; tibia 160–181; tarsus 148–170].
Gnathosoma (Fig.
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [447–476 total; trochanter 59–63; basifemur 80–88; telofemur 61–68; genu 79–88; tibia 85–95; tarsus 80–92]. Leg-II [479–526 total; trochanter 54–67; basifemur 82–89; telofemur 60–72; genu 80–90; tibia 94–105; tarsus 105–114]. Leg-III [544–624 total; trochanter 56–66; basifemur 79–102; telofemur 65–74; genu 95–110; tibia 119–138; tarsus 120–137]. Leg-IV [743–857 total; trochanter 85–110; basifemur 107–125; telofemur 113–130; genu 134–160; tibia 152–177; tarsus 145–158].
Same as genus.
Same as genus.
Members of this genus, unlike
Same as subfamily.
Holotype (1♂) deposited at the
Gnathosoma — Subcapitulum [154–173 ventral length; 96–108 dorsal length; 90–105 tall] elliptic to ovoid with short rostrum. Chelicerae [133–152 long] unmodified with slightly curved fangs [29–32 long]. Pedipalp [181–202 long] unmodified. Trochanter [25–30 long; 30–35 wide]. Femur [49–58 long; 38–42 wide]. Genu [38–42 long; 32–35 wide]. Tibia [45–52 long; 22–25 wide]. Tarsus [19–23 long; 9–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless, or with same color as dorsal plate. Total leg and podomere lengths as follows: Leg-I [428–477 total; trochanter 48–55; basifemur 72–85; telofemur 60–69; genu 78–90; tibia 83–95; tarsus 79–92]. Leg-II [453–530 total; trochanter 54–62; basifemur 74–87; telofemur 58–68; genu 83–96; tibia 96–110; tarsus 99–113]. Leg-III [440–625 total; trochanter 55–65; basifemur 76–88; telofemur 64–76; genu 106–117; tibia 120–137; tarsus 131–148]. Leg-IV [677–843 total; trochanter 87–97; basifemur 106–120; telofemur 111–122; genu 146–160; tibia 160–173; tarsus 147–180].
Gnathosoma — Subcapitulum [138–164 ventral length; 88–105 dorsal length; 83–93 tall]. Chelicerae [120–145 long]. Fangs [27–30 long]. Pedipalp [181–206 long]. Trochanter [24–32 long; 28–33 wide]. Femur [48–59 long; 35–40 wide]. Genu [38–46 long; 29–34 wide]. Tibia [43–54 long; 19–25 wide]. Tarsus [16–22 long; 9–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [435–483 total; trochanter 53–63; basifemur 75–84; telofemur 57–69; genu 78–89; tibia 82–93; tarsus 80–90]. Leg-II [458–518 total; trochanter 52–64; basifemur 75–87; telofemur 59–69; genu 79–90; tibia 92–104; tarsus 96–109]. Leg-III [530–599 total; trochanter 54–62; basifemur 75–88; telofemur 63–72; genu 97–111; tibia 114–133; tarsus 124–137]. Leg-IV [722–813 total; trochanter 81–95; basifemur 102–122; telofemur 103–118; genu 130–159; tibia 150–167; tarsus 145–158].
Abundant throughout North America, ranging from the Pacific Northwest to the southwestern United States (and potentially into northern Mexico), and east into the western Great Plains.
Commonly colorless or orange in the southwestern United States; red, pink, or orange–red in the northwest, Rocky Mountains, and western Great Plains; and uncommonly red–violent in the northwest, Rocky Mountains, and western Great Plains.
Syntypes (1♀, 1♂) deposited at the
Gnathosoma — Subcapitulum [151–190 ventral length; 90–114 dorsal length; 84–115 tall] elliptical to ovoid with short rostrum. Chelicerae [133–170 long] unmodified with lightly curved fangs [28–35 long]. Pedipalp [169–211 long] unmodified. Trochanter [23–32 long; 28–37 wide]. Femur [46–62 long; 33–45 wide]. Genu [33–42 long; 28–36 wide]. Tibia [42–53 long; 19–26 wide]. Tarsus [18–23 long; 9–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless, or with same color as dorsal plate. Total leg and podomere lengths as follows: Leg-I [401–497 total; trochanter 50–61; basifemur 74–85; telofemur 55–72; genu 72–96; tibia 75–97; tarsus 78–97]. Leg-II [417–564 total; trochanter 51–63; basifemur 71–92; telofemur 57–72; genu 75–100; tibia 92–118; tarsus 96–120]. Leg-III [513–664 total; trochanter 55–68; basifemur 71–96; telofemur 58–82; genu 91–124; tibia 112–147; tarsus 124–157]. Leg-IV [726–911 total; trochanter 85–105; basifemur 103–132; telofemur 99–138; genu 134–174; tibia 145–177; tarsus 148–185].
Gnathosoma — Subcapitulum [128–155 ventral length; 83–96 dorsal length; 78–95 tall]. Chelicerae [115–145 long]. Fangs [25–29 long]. Pedipalp [156–190 long]. Trochanter [22–28 long; 28–33 wide]. Femur [42–55 long; 32–42 wide]. Genu [32–41 long; width 25–32 wide]. Tibia [43–52 long; 19–23 wide]. Tarsus [16–21 long; 9–11 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [402–452 total; trochanter 49–59; basifemur 67–80; telofemur 53–63; genu 70–82; tibia 75–88; tarsus 78–88]. Leg-II [421–488 total; trochanter 51–61; basifemur 68–81; telofemur 51–63; genu 73–86; tibia 84–96; tarsus 91–105]. Leg-III [501–552 total; trochanter 52–61; basifemur 72–82; telofemur 59–68; genu 89–100; tibia 105–119; tarsus 118–130]. Leg-IV [664–746 total; trochanter 79–90; basifemur 95–106; telofemur 92–108; genu 124–144; tibia 130–155; tarsus 129–150].
Abundant throughout the majority of North America. Unreported in Washington and northern Oregon.
Commonly orange and uncommonly violet in the southwestern United States; commonly violet or blue and uncommonly red–violet in the Rocky Mountains and Great Plains; commonly violet or blue east of the Great Plains.
Holotype (1♀) and allotype (1♂) deposited at the
Gnathosoma — Subcapitulum [174 ventral length; 104 dorsal length; 90 tall] elliptical to ovoid with short rostrum and colorless. Chelicerae [144 long] unmodified with lightly curved fangs [32 long]. Pedipalp [190 long] unmodified. Trochanter [28 long; 29 wide]. Femur [53 long; 42 wide]. Genu [39 long; 32 wide]. Tibia [50 long; 23 wide]. Tarsus [19 long; 10 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [480 total; trochanter 62; basifemur 80; telofemur 64; genu 91; tibia 92; tarsus 90]. Leg-II [519 total; trochanter 63; basifemur 83; telofemur 69; genu 94; tibia 104; tarsus 106]. Leg-III [615 total; trochanter 63; basifemur 85; telofemur 72; genu 115; tibia 133; tarsus 145]. Leg-IV [821 total; trochanter 93; basifemur 112; telofemur 122; genu 161; tibia 178; tarsus 155].
Gnathosoma — Subcapitulum [139 ventral length; 90 dorsal length; 83 tall]. Chelicerae [125 long]. Fangs [29 long]. Pedipalp [179 long]. Trochanter [26 long; 29 wide]. Femur [48 long; 35 wide]. Genu [40 long; width 29 wide]. Tibia [44 long; 23 wide]. Tarsus [20 long; 10 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [413 total; trochanter 51; basifemur 69; telofemur 61; genu 73; tibia 79; tarsus 78]. Leg-II [462 total; trochanter 60; basifemur 75; telofemur 59; genu 80; tibia 94; tarsus 93]. Leg-III [517 total; trochanter 56; basifemur 73; telofemur 65; genu 95; tibia 111; tarsus 116]. Leg-IV [688 total; trochanter 76; basifemur 97; telofemur 97; genu 132; tibia 146; tarsus 138].
Specific epithet
Known from only two counties (Los Angeles and Sierra) in California.
Holotype (1♀), allotype (1♂), and three paratypes (3♂) deposited at the
Gnathosoma — Subcapitulum [153–155 ventral length; 117–133 dorsal length; 88–97 tall] ovoid with short rostrum. Chelicerae [148–156 long] unmodified with lightly curved fangs [28–29 long]. Pedipalp [177–187 long] unmodified and violet. Trochanter [27–30 long; 26–29 wide]. Femur [46–51 long; 35–38 wide]. Genu [38–42 long; 27–28 wide]. Tibia [44–49 long; 19–20 wide]. Tarsus [18–19 long; 10–11 wide].
Dorsum (Fig.
Venter (Fig.
Legs — violet. Total leg and podomere lengths as follows: Leg-I [464–466 total; trochanter 57–58; basifemur 81–82; telofemur 65–68; genu 83–84; tibia 88–90; tarsus 86–87]. Leg-II [489–490 total; trochanter 54–55; basifemur 81–83; telofemur 64–66; genu 86–87; tibia 97–101; tarsus 102–105]. Leg-III [559–564 total; trochanter 57–58; basifemur 77–85; telofemur 73–76; genu 102–105; tibia 116–117; tarsus 126–130]. Leg-IV [760–767 total; trochanter 86–87; basifemur 107–108; telofemur 108–109; genu 145–146; tibia 158–159; tarsus 152–159].
Gnathosoma — Subcapitulum [132–143 ventral length; 85–90 dorsal length; 81–86 tall]. Chelicerae [107–115 long]. Fangs [25–28 long]. Pedipalp [170–181 long]. Trochanter [25–27 long; 28–30 wide]. Femur [45–52 long; 33–35 wide]. Genu [38–39 long; width 27–29 wide]. Tibia [45–50 long; 18–21 wide]. Tarsus [14–17 long; 8–11 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [440–454 total; trochanter 53–58; basifemur 76–80; telofemur 58–67; genu 75–80; tibia 84–89; tarsus 82–90]. Leg-II [464–478 total; trochanter 52–57; basifemur 75–80; telofemur 58–62; genu 78–86; tibia 94–97; tarsus 99–103]. Leg-III [512–535 total; trochanter 49–55; basifemur 74–83; telofemur 62–69; genu 93–96; tibia 106–116; tarsus 110–125]. Leg-IV [699–726 total; trochanter 77–87; basifemur 101–110; telofemur 99–108; genu 130–133; tibia 144–148; tarsus 133–147].
Specific epithet
Reported from only two counties (Polk and Montgomery) in Arkansas.
Holotype (1♀), allotype (1♂), and eight paratypes (4♀, 4♂) deposited at
These mites differ from all others in the complex in having large medial pores on the dorsal plate surrounded by a ring of smaller pores (all pores uniform in other species). Males also have a “bleached” or colorless area posterior to the coxal plate that is colored in other members of the complex.
Gnathosoma — Subcapitulum [165–175 ventral length; 99–106 dorsal length; 90–100 tall] ovoid with short rostrum. Chelicerae [139–150 long] unmodified with lightly curved fangs [29–32 long]. Pedipalp [192–205 long] unmodified. Trochanter [25–28 long; 29–32 wide]. Femur [54–57 long; 37–40 wide]. Genu [40–46 long; 29–33 wide]. Tibia [51–55 long; 20–23 wide]. Tarsus [19–21 long; 10–11 wide].
Dorsum (Fig.
Venter (Fig.
Legs — orange and restricted violet to blue. Total leg and podomere lengths as follows: Leg-I [473–524 total; trochanter 60–62; basifemur 83–93; telofemur 65–76; genu 86–95; tibia 92–105; tarsus 83–95]. Leg-II [501–552 total; trochanter 54–63; basifemur 83–93; telofemur 65–72; genu 88–99; tibia 101–111; tarsus 102–115]. Leg-III [586–635 total; trochanter 61–65; basifemur 89–100; telofemur 70–80; genu 105–113; tibia 122–137; tarsus 132–144]. Leg-IV [805–876 total; trochanter 93–109; basifemur 115–132; telofemur 115–125; genu 151–167; tibia 167–180; tarsus 158–177].
Gnathosoma — Subcapitulum [150–160 ventral length; 95–106 dorsal length; 86–95 tall]. Chelicerae 127–139 long]. Fangs [26–29 long]. Pedipalp [180–195long]. Trochanter [25–27 long; 27–30 wide]. Femur [50–55 long; 34–37 wide]. Genu [38–41 long; width 26–29 wide]. Tibia [49–52 long; 19–22 wide]. Tarsus [17–20 long; 9–11 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [444–508 total; trochanter 55–62; basifemur 75–89; telofemur 63–73; genu 80–91; tibia 85–99; tarsus 84–96]. Leg-II [474–533 total; trochanter 60–64; basifemur 77–90; telofemur 61–71; genu 82–93; tibia 92–106; tarsus 99–113]. Leg-III [537–598 total; trochanter 57–64; basifemur 80–92; telofemur 65–73; genu 96–108; tibia 113–128; tarsus 121–137]. Leg-IV [721–778 total; trochanter 88–99; basifemur 96–117; telofemur 102–113; genu 135–151; tibia 147–168; tarsus 142–156].
Specific epithet
Eastern United States east of the Mississippi River with southern limits in Florida.
As it is likely that this species represents a cryptic species complex, measurements were only included from specimens exhibiting less than 2% COI divergence within the clade (those highlighted in red in Fig.
Holotype (1♀), allotype (1♂) and ten paratypes (5♀, 5♂) deposited at Canadian National Collection; eleven paratypes (7♀, 4♂) at
These mites have violet to blue coloration over the majority of their anterio-lateral platelets while the rest of the complex have coloration restricted to the posterior half of the platelet.
Gnathosoma — Subcapitulum [143–165 ventral length; 90–105 dorsal length; 84–95 tall] ovoid with short rostrum. Chelicerae [119–136 long] unmodified with lightly curved fangs [24–32 long]. Pedipalp [167–191 long] unmodified. Trochanter [23–30 long; 29–31 wide]. Femur [47–53 long; 33–38 wide]. Genu [37–42 long; 27–30 wide]. Tibia [40–53 long; 17–22 wide]. Tarsus [15–20 long; 9–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — violet to blue and orange. Total leg and podomere lengths as follows: Leg-I [449–485 total; trochanter 54–62; basifemur 77–83; telofemur 62–70; genu 80–90; tibia 89–99; tarsus 80–90]. Leg-II [471–510 total; trochanter 54–60; basifemur 74–84; telofemur 61–66; genu 82–94; tibia 98–107; tarsus 87–109]. Leg-III [548–612 total; trochanter 55–64; basifemur 79–91; telofemur 66–74; genu 96–114; tibia 116–137; tarsus 119–141]. Leg-IV [737–825 total; trochanter 79–99; basifemur 103–123; telofemur 103–121; genu 138–154; tibia 154–169; tarsus 147–167].
Gnathosoma — Subcapitulum [133–144 ventral length; 83–90 dorsal length; 72–84 tall]. Chelicerae [110–120 long]. Fangs [25–30 long]. Pedipalp [168–183 long]. Trochanter [22–25 long; 25–29 wide]. Femur [45–50 long; 32–36 wide]. Genu [36–40 long; width 24–30 wide]. Tibia [44–52 long; 18–20 wide]. Tarsus [16–20 long; 8–11 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [419–451 total; trochanter 45–56; basifemur 70–77; telofemur 58–68; genu 74–82; tibia 81–90; tarsus 79–84]. Leg-II [429–472 total; trochanter 47–52; basifemur 69–77; telofemur 56–63; genu 76–86; tibia 86–98; tarsus 93–99]. Leg-III [491–540 total; trochanter 49–53; basifemur 70–86; telofemur 59–66; genu 89–98; tibia 107–120; tarsus 114–124]. Leg-IV [665–739 total; trochanter 74–90; basifemur 95–109; telofemur 95–108; genu 128–138; tibia 139–150; tarsus 131–145].
Specific epithet after Samuel Benjamin Harris, the American author, philosopher, and co-founder of Project Reason. Sam Harris, more than any speaker and author, has challenged my (JCO) views and assumptions and kept me on my toes.
Eastern United States east of the Mississippi river, with southern limits in Florida.
Holotype (1♀), allotype (1♂), and six paratypes (3♀, 3♂) deposited at
Both
Gnathosoma — Subcapitulum [139–152 ventral length; 85–97 dorsal length; 85–91 tall] ovoid with short rostrum. Chelicerae [117–126 long] unmodified with lightly curved fangs [24–28 long]. Pedipalp [168–189 long] unmodified. Trochanter [23–27 long; 28–31 wide]. Femur [42–52 long; 33–35 wide]. Genu [35–41 long; 25–32 wide]. Tibia [45–52 long; 17–22 wide]. Tarsus [18–20 long; 8–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — violet to blue and orange. Total leg and podomere lengths as follows: Leg-I [431–463 total; trochanter 48–58; basifemur 70–78; telofemur 59–65; genu 77–84; tibia 85–93; tarsus 81–88]. Leg-II [455–487 total; trochanter 51–56; basifemur 72–79; telofemur 57–64; genu 80–88; tibia 92–102; tarsus 98–109]. Leg-III [538–572 total; trochanter 54–59; basifemur 73–83; telofemur 62–67; genu 95–106; tibia 114–127; tarsus 128–134]. Leg-IV [641–768 total; trochanter 84–89; basifemur 96–115; telofemur 102–110; genu 137–144; tibia 147–163; tarsus 142–158].
Gnathosoma — Subcapitulum [125–134 ventral length; 80–86 dorsal length; 74–83 tall]. Chelicerae [100–115 long]. Fangs [24–28 long]. Pedipalp [164–179 long]. Trochanter [22–24 long; 26–29 wide]. Femur [44–50 long; 30–35 wide]. Genu [36–43 long; width 25–28 wide]. Tibia [44–49 long; 17–20 wide]. Tarsus [17–19 long; 9–10 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [414–434 total; trochanter 47–54; basifemur 67–73; telofemur 55–62; genu 72–79; tibia 81–85; tarsus 79–85]. Leg-II [432–450 total; trochanter 48–54; basifemur 66–72; telofemur 54–61; genu 73–80; tibia 88–91; tarsus 96–99]. Leg-III [478–525 total; trochanter 49–58; basifemur 66–76; telofemur 58–64; genu 83–93; tibia 102–114; tarsus 114–126]. Leg-IV [658–685 total; trochanter 76–86; basifemur 85–103; telofemur 90–100; genu 124–130; tibia 140–141; tarsus 133–140].
Specific epithet
Eastern United States east of the Mississippi River, with southern limits in Florida.
Holotype (1♀), allotype (1♂), and six paratypes (3♀, 3♂) deposited at
Both
Gnathosoma — Subcapitulum [160–168 ventral length; 102–105 dorsal length; 91–95 tall] ovoid with short rostrum. Chelicerae [136–141 long] unmodified with lightly curved fangs [29–30 long]. Pedipalp [188–193 long] unmodified. Trochanter [26–29 long; 28–32 wide]. Femur [50–54 long; 35–37 wide]. Genu [39–41 long; 30–33 wide]. Tibia [51–52 long; 19–22 wide]. Tarsus [19–20(–21) long; 9–11 wide].
Dorsum (Fig.
Venter (Fig.
Legs — violet to blue. Total leg and podomere lengths as follows: Leg-I [487–500 total; trochanter 57–63; basifemur 84–86; telofemur 67–73; genu 90–94; tibia 94–99; tarsus 87–94]. Leg-II [532–548 total; trochanter 58–65; basifemur 84–89; telofemur 67–72; genu 94–99; tibia 107–113; tarsus 110–116]. Leg-III [599–629 total; trochanter 63–68; basifemur 88–97; telofemur 73–76; genu 107–117; tibia 128–138; tarsus 134–140]. Leg-IV [830–861 total; trochanter 83–104; basifemur 113–130; telofemur 119–130; genu 156–164; tibia 172–181; tarsus 164–175].
Gnathosoma — Subcapitulum [143–156 ventral length; 89–97 dorsal length; 81–91 tall]. Chelicerae [113–129 long]. Fangs [26–30 long]. Pedipalp [180–195 long]. Trochanter [24–30 long; 26–29 wide]. Femur [50–53 long; 33–36 wide]. Genu [38–43 long; width 27–29 wide]. Tibia [47–52 long; 19–22 wide]. Tarsus [17–20 long; 9–10 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [452–497 total; trochanter 52–63; basifemur 78–87; telofemur 63–72; genu 84–92; tibia 89–99; tarsus 84–93]. Leg-II [486–519 total; trochanter 53–61; basifemur 77–85; telofemur 59–70; genu 87–91; tibia 102–107; tarsus 101–112]. Leg-III [551–588 total; trochanter 54–60; basifemur 81–90; telofemur 66–73; genu 100–107; tibia 118–129; tarsus 126–134]. Leg-IV [752–796 total; trochanter 85–94; basifemur 99–120; telofemur 107–117; genu 143–146; tibia 158–163; tarsus 148–158].
Eastern United States east of the Mississippi River, with southern limits in Florida.
Specific epithet
Holotype (1♀) deposited at
Resembling most
Gnathosoma — Subcapitulum [180–199 ventral length; 108–121 dorsal length; 110–124 tall] ovoid with short rostrum and colorless. Chelicerae [148–173 long] unmodified with lightly curved fangs [30–33 long]. Pedipalp [209–236 long] unmodified. Trochanter [29–37 long; 31–33 wide]. Femur [54–59 long; 38–45 wide]. Genu [47–60 long; 32–36 wide]. Tibia [53–59 long; 21–24 wide]. Tarsus [19–23 long; 9–13 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [542–604 total; trochanter 68–75; basifemur 91–108; telofemur 75–83; genu 95–111; tibia 108–122; tarsus 101–112]. Leg-II [548–611 total; trochanter 63–72; basifemur 95–108; telofemur 70–83; genu 93–103; tibia 107–123; tarsus 110–127]. Leg-III [595–683 total; trochanter 66–73; basifemur 99–108; telofemur 76–84; genu 103–124; tibia 127–150; tarsus 124–153]. Leg-IV [854–987 total; trochanter 94–112; basifemur 125–152; telofemur 134–154; genu 170–192; tibia 171–216; tarsus 154–181].
Gnathosoma — Subcapitulum [164–178 ventral length; 97–109 dorsal length; 96–114 tall]. Chelicerae [132–152 long]. Fangs [28–33 long]. Pedipalp [202–222 long]. Trochanter [28–33 long; 28–33 wide]. Femur [50–58 long; 38–42 wide]. Genu [50–55 long; width 30–35 wide]. Tibia [53–58 long; 20–22 wide]. Tarsus [18–23 long; 11–13 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [501–560 total; trochanter 57–64; basifemur 85–99; telofemur 70–80; genu 90–102; tibia 101–113; tarsus 95–108]. Leg-II [508–567 total; trochanter 58–67; basifemur 88–96; telofemur 67–74; genu 83–99; tibia 101–117; tarsus 105–119]. Leg-III [554–615 total; trochanter 59–63; basifemur 83–98; telofemur 70–76; genu 97–115; tibia 117–138; tarsus 119–132]. Leg-IV [526–882 total; trochanter 79–103; basifemur 116–130; telofemur 121–135; genu 157–175; tibia 171–197; tarsus 149–166].
Western North America within and west of the Rocky Mountains. California (
Having examined the type material, we suggest that
Holotype (1♀) deposited at
These are the largest known testudacarines and also differ further from their complex in having a smaller, more rounded anterio-medial platelet.
Gnathosoma — Subcapitulum [245 ventral length; 133 dorsal length; 130 tall] ovoid with short rostrum. Chelicerae [200 long] unmodified with lightly curved fangs [40 long]. Pedipalp [259 long] unmodified. Trochanter [38 long; 43 wide]. Femur [70 long; 50 wide]. Genu [63 long; 40 wide]. Tibia [63 long; 28 wide]. Tarsus [25 long; 15 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [620 total; trochanter 70; basifemur 119; telofemur 89; genu 101; tibia 119; tarsus 121]. Leg-II [681 total; trochanter 77; basifemur 113; telofemur 90; genu 117; tibia 145; tarsus 140]. Leg-III [756 total; trochanter 70; basifemur 116; telofemur 93; genu 144; tibia 165; tarsus 167]. Leg-IV [1081 total; trochanter 136; basifemur 148; telofemur 159; genu 207; tibia 224; tarsus 208].
Specific epithet
As it is likely that this species represents a cryptic species complex (with 4.5% COI divergence between the two available specimens), measurements were included from only one specimen (the other highlighted in red in Fig.
Only one specimen know from Douglas County, Oregon.
Holotype (1♀) and allotype (1♂) deposited at
These mites differ from the rest of the complex in having a dorsally “covered” gnathosomal bay (short doral gnathosomal bay length) and an elongate gnathosoma with a long rostrum that exteneds below the gnathosoma ventral surface.
Gnathosoma (Fig.
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [566–594 total; trochanter 68–74; basifemur 104–106; telofemur 81–85; genu 108–112; tibia 109–117; tarsus 95–102]. Leg-II [614–645 total; trochanter 75–78; basifemur 102–108; telofemur 77–79; genu 111–116; tibia 124–130; tarsus 125–135]. Leg-III [714–753 total; trochanter 74–79; basifemur 109–119; telofemur 88–94; genu 136–139; tibia 154–160; tarsus 152–161]. Leg-IV [952–961 total; trochanter 105–109; basifemur 142–144; telofemur 138–139; genu 191–192; tibia 199–201; tarsus 175–178].
Gnathosoma (Fig.
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [593–606 total; trochanter 73–79; basifemur 111–114; telofemur 78–83; genu 107–111; tibia 113–118; tarsus 105–106]. Leg-II [635–645 total; trochanter 74–79; basifemur 102–104; telofemur 84–85; genu 115–120; tibia 129–132; tarsus 124–130]. Leg-III [724–726 total; trochanter 77–78; basifemur 109–116; telofemur 87–91; genu 136–137; tibia 155–156; tarsus 152–155]. Leg-IV [905–964 total; trochanter 107–118; basifemur 139–140; telofemur 129–142; genu 183–191; tibia 179–198; tarsus 167–176].
Specific epithet
Humbolt County, California and Curry County, Oregon.
Holotype (1♀), allotype (1♂), and eight paratypes (4♀, 4♂) deposited at
Resembling most
Gnathosoma — Subcapitulum [217–245 ventral length; 137–145 dorsal length; 125–152 tall] elliptical to ovoid with short rostrum. Chelicerae [190–205 long] unmodified with lightly curved fangs [40–45 long]; fangs characteristically large. Pedipalp [250–272 long] unmodified. Trochanter [37–45 long; 39–46 wide]. Femur [72–80 long; 52–61 wide]. Genu [55–61 long; 41–49 wide]. Tibia [57–66 long; 23–26 wide]. Tarsus [21–26 long; 11–13 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [624–660 total; trochanter 74–81; basifemur 110–121; telofemur 84–95; genu 117–125; tibia 123–130; tarsus 108–116]. Leg-II [658–718 total; trochanter 75–87; basifemur 111–123; telofemur 83–95; genu 115–130; tibia 133–149; tarsus 133–145]. Leg-III [755–820 total; trochanter 78–89; basifemur 112–131; telofemur 88–102; genu 139–155; tibia 162–184; tarsus 165–182]. Leg-IV [1017–1058 total; trochanter 117–132; basifemur 139–150; telofemur 140–153; genu 197–210; tibia 212–228; tarsus 194–205].
Gnathosoma — Subcapitulum [205–232 ventral length; 125–145 dorsal length; 124–133 tall]. Chelicerae [178–202 long]. Fangs [39–42 long]. Pedipalp [250–279 long]. Trochanter [37–42 long; 38–43 wide]. Femur [71–81 long; 52–60 wide]. Genu [55–65 long; width 40–47 wide]. Tibia [59–69 long; 23–25 wide]. Tarsus [21–27 long; 11–15 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [617–679 total; trochanter 73–80; basifemur 110–122; telofemur 84–97; genu 116–129; tibia 118–138; tarsus 107–123]. Leg-II [664–743 total; trochanter 74–84; basifemur 110–125; telofemur 85–103; genu 118–137; tibia 134–157; tarsus 131–148]. Leg-III [753–841 total; trochanter 76–83; basifemur 111–133; telofemur 89–106; genu 138–158; tibia 163–186; tarsus 155–186]. Leg-IV [952–1098 total; trochanter 111–131; basifemur 136–154; telofemur 139–161; genu 189–217; tibia 189–238; tarsus 176–210].
Specific epithet
British Columbia, Canada.
Holotype (1♀) and allotype (1♂) deposited at
These mites are smaller and more colorful than other species in the complex and therefore resemble most the
Gnathosoma — Subcapitulum [176–188 ventral length; 105–107 dorsal length; 100–103 tall] elliptical ovoid with short rostrum and colorless. Chelicerae [145–152 long] unmodified with lightly curved fangs [28–30 long]. Pedipalp [202–212 long] unmodified. Trochanter [30–31 long; 28–30 wide]. Femur [56–58 long; 40–41 wide]. Genu [43–47 long; 33–35 wide]. Tibia [52–55 long; 20–23 wide]. Tarsus [20–22 long; 9–10 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [503–542 total; trochanter 65–68; basifemur 88–93; telofemur 70–81; genu 93–103; tibia 96–102; tarsus 89–97]. Leg-II [510–577 total; trochanter 52–74; basifemur 82–94; telofemur 66–78; genu 94–102; tibia 103–118; tarsus 106–112]. Leg-III [610–657 total; trochanter 64–71; basifemur 92–99; telofemur 73–82; genu 110–122; tibia 127–146; tarsus 137–141]. Leg-IV [843–914 total; trochanter 96–101; basifemur 116–125; telofemur 117–133; genu 166–185; tibia 181–195; tarsus 168–178].
Gnathosoma — Subcapitulum [160–170 ventral length; 98–109 dorsal length; 81–92 tall]. Chelicerae [132–140 long]. Fangs [27–29 long]. Pedipalp [184–190 long]. Trochanter [23–26 long; 29–30 wide]. Femur [50–51 long; 36–38 wide]. Genu [41–42 long; width 29–30 wide]. Tibia [49–52 long; 19–20 wide]. Tarsus [17–21 long; 8–9 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [472 total; trochanter 59–60; basifemur 83–91; telofemur 66–71; genu 88–91; tibia 91–97; tarsus 83–88]. Leg-II [496–515 total; trochanter 61–66; basifemur 84–87; telofemur 65–69; genu 85–97; tibia 96–108; tarsus 102–107]. Leg-III [554–593 total; trochanter 62–69; basifemur 84–89; telofemur 65–74; genu 100–110; tibia 117–126; tarsus 125–136]. Leg-IV [784–822 total; trochanter 80–95; basifemur 109–116; telofemur 114–115; genu 155–162; tibia 164–174; tarsus 155–162].
Specific epithet
Reported from only two counties (Mendocino and Nevada) in California.
Combined molecular, distributional, and morphological data support three distinct clades within the
Holotype (1♀), allotype (1♂), and four paratypes (3♀, 1♂) deposited at
Since morphological variation is limited, a combination of morphology and distribution is best used to diagnose members of the complex. These mites occur in Washington within and east of the Cascade Mountains, while
Gnathosoma — Subcapitulum [174–175 ventral length; 110–118 dorsal length; 102–122 tall] ovoid with short rostrum. Chelicerae [140–163 long] unmodified with lightly curved fangs [33–37 long]. Pedipalp [221–248 long] unmodified. Trochanter [28–38 long; 32–36 wide]. Femur [60–64 long; 45–50 wide]. Genu [51–59 long; 37–43 wide]. Tibia [61–69 long; 24–28 wide]. Tarsus [20–25 long; 10–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [561–614 total; trochanter 63–66; basifemur 101–114; telofemur 79–93; genu 103–116; tibia 110–127; tarsus 96–109]. Leg-II [559–623 total; trochanter 56–65; basifemur 96–120; telofemur 75–88; genu 103–116; tibia 120–128; tarsus 107–120]. Leg-III [630–703 total; trochanter 60–80; basifemur 97–116; telofemur 79–96; genu 116–139; tibia 136–152; tarsus 129–140]. Leg-IV [863–920 total; trochanter 98–109; basifemur 126–140; telofemur 130–140; genu 172–189; tibia 183–194; tarsus 152–167].
Gnathosoma — Subcapitulum [148–160 ventral length; 98–108 dorsal length; 95–100 tall]. Chelicerae [135–140 long]. Fangs [30–31 long]. Pedipalp [208–215 long]. Trochanter [30–31 long; 28–33 wide]. Femur [53–60 long; 40–44 wide]. Genu [47–52 long; width 33–35 wide]. Tibia [55–61 long; 23–25 wide]. Tarsus [20–21 long; 10–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [531–558 total; trochanter 54–65; basifemur 95–99; telofemur 75–80; genu 98–104; tibia 105–111; tarsus 100–105]. Leg-II [549–572 total; trochanter 59–65; basifemur 93–101; telofemur 73–79; genu 98–105; tibia 112–120; tarsus 100–109]. Leg-III [577–610 total; trochanter 64–69; basifemur 93–105; telofemur 75–84; genu 106–119; tibia 122–135; tarsus 117–131]. Leg-IV [765–812 total; trochanter 91–100; basifemur 110–120; telofemur 115–121; genu 153–171; tibia 152–168; tarsus 140–152].
Within and east of the Cascade Mountains, Washington.
Holotype (1♂) deposited at
Since morphological variation is limited, a combination of morphology and distribution is best used to diagnosed members of the complex. These mites occur in the Olympic Mountains, while
Gnathosoma — Subcapitulum [173 ventral length; 108 dorsal length; 105 tall] ovoid with short rostrum. Chelicerae [150 long] unmodified with lightly curved fangs [33-37 long]. Pedipalp [249 long] unmodified. Trochanter [35 long; 34 wide]. Femur [60 long; 48 wide]. Genu [56 long; 40 wide]. Tibia [75 long; 25 wide]. Tarsus [23 long; 12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [603 total; trochanter 61; basifemur 103; telofemur 89; genu 116; tibia 124; tarsus 108]. Leg-II [610 total; trochanter 63; basifemur 101; telofemur 85; genu 115; tibia 126; tarsus 120]. Leg-III [674 total; trochanter 63; basifemur 110; telofemur 86; genu 130; tibia 145; tarsus 137]. Leg-IV [870 total; trochanter 87; basifemur 123; telofemur 130; genu 179; tibia 189; tarsus 160].
Specific epithet
One specimen found in Mason County in the Olympic Mountains, Washington.
Holotype (1♀), allotype (1♂), and ten paratypes (6♀, 4♂) deposited at
Since morphological variation is limited, a combination of morphology and distribution is best used to diagnosed members of the complex. These mites occur along the western coast of Washington, Oregon, California, and British Columbia, while
Gnathosoma — Subcapitulum [192–208 ventral length; 116–132 dorsal length; 120–134 tall] ovoid with short rostrum. Chelicerae [149–166 long] unmodified with lightly curved fangs [33–36 long]. Pedipalp [231–242 long] unmodified. Trochanter [31–36 long; 30–33 wide]. Femur [58–63 long; 45–49 wide]. Genu [53–59 long; 36–38 wide]. Tibia [64–69 long; 22–25 wide]. Tarsus [20–25 long; 11–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — colorless. Total leg and podomere lengths as follows: Leg-I [623–676 total; trochanter 72–85; basifemur 106–115; telofemur 85–94; genu 114–129; tibia 123–137; tarsus 109–120]. Leg-II [642–689 total; trochanter 75–80; basifemur 108–117; telofemur 88–94; genu 115–135; tibia 131–152; tarsus 119–133]. Leg-III [710–777 total; trochanter 70–80; basifemur 106–126; telofemur 92–100; genu 129–151; tibia 151–172; tarsus 146–161]. Leg-IV [941–1001 total; trochanter 106–125; basifemur 135–150; telofemur 139–146; genu 188–199; tibia 197–215; tarsus 160–178].
Gnathosoma — Subcapitulum [153–177 ventral length; 99–118 dorsal length; 98–110 tall]. Chelicerae [123–148 long]. Fangs [28–32 long]. Pedipalp [201–231 long]. Trochanter [29–33 long; 27–30 wide]. Femur [51–55 long; 37–46 wide]. Genu [44–55 long; width 32–37 wide]. Tibia [54–66 long; 20–23 wide]. Tarsus [19–22 long; 10–12 wide].
Dorsum (Fig.
Venter (Fig.
Legs — total leg and podomere lengths as follows: Leg-I [526–617 total; trochanter 57–69; basifemur 90–103; telofemur 75–90; genu 100–116; tibia 105–124; tarsus 97–116]. Leg-II [536–629 total; trochanter 59–70; basifemur 87–106; telofemur 73–86; genu 100–117; tibia 106–134; tarsus 104–123]. Leg-III [589–691 total; trochanter 60–73; basifemur 89–111; telofemur 71–90; genu 115–130; tibia 123–151; tarsus 128–147]. Leg-IV [810–878 total; trochanter 92–101; basifemur 112–125; telofemur 114–127; genu 144–177; tibia 158–188; tarsus 143–168]
Specific epithet
West coast of British Columbia, Washington, Oregon, and California.
Holotype (1♀) at Naturhistorisches Museum Basel, Switzerland.
India (
The types were not examined for this publication.
Holotype (1♂) at Taiji Imamura Collection at Ibaraki Nature Museum, Japan.
Holotype loans are not available from Ibaraki Nature Museum. The museum provided a low-magnification photograph through e-mail, though permission to print the photograph was not obtained.
These mites differ from all other
Takékura, Japan (
It is reasonable to assume
The types were not examined for this publication.
Holotype (1♀) and allotype (1♂?) at Taiji Imamura Collection at Ibaraki Nature Museum, Japan.
Holotype loans are not available from Ibaraki Nature Museum. The museum provided a low-magnification photograph through e-mail, though permission to print the photograph was not obtained.
These mites differ from all other Asian
Throughout Honshu, Japan (
A drawing of the “male” dorsum is left out of the
The types were not examined for this publication; contact with the authors was attempted but unsuccessful and the the types were not examined.
Institute of Entomology, Guizhou University.
These mites can differ from all other
Mt. Fanjing (
1 | Pedipalp four-segmented, anterior tip of coxa-I with projection; California |
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– | Pedipalp five-segmented, anterior tip of coxa-I without projection |
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2 | Body elongate to rectangular | ( |
– | Body oval |
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3 | Lateral platelets robust; lateral-platelet-4 large and in direct or near direct contact with lateral-platelet-2 |
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– | Lateral-platelet-4 small, gap present between it and lateral-platlet-2; widespread along west coast of N. America |
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4 | Distribution restricted to the Olympic Mountains |
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– | Distribution within and east of the Cascade Mountains |
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5 | Body large (>700 µm female and >650 male dorsal length; if smaller, found in Asia), dull coloration common; within and west of the Rocky Mountains or Asia | (Asian species and |
– | Body small (<700 µm female and <650 male dorsal length), bright coloration (orange, red, violet, blue) common; present throughout North America | ( |
6 | Gnathosomal bay “covered” (short dorsal gnathosomal bay length), gnathosoma elongate with long rostrum that extends below ventral surface of gnathosoma |
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– | Without these characters |
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7 | Body very large (>900 µm dorsal length, female) with small, square (unusual for complex) anterio-medial platelet (male unknown) |
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– | Body not this large (<900 µm dorsal length, female and male) often with wide, rectangular or trapezoidal anterio-medial platelet |
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8 | Anterior-medial platelet compact and pentagonal, suture lines between second and third coxae absent; India, Java, Bhutan |
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– | Anterior-medial platelet wider, more trapezoidal in shape, suture lines between second and third coxae present, but incomplete |
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9 | Body small (<650 µm dorsal length female and <450 µm dorsal length male); Japan |
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– | Body larger (>700 µm dorsal length female and >500 µm dorsal length male) |
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10 | Body <770 µm dorsal length female and <660 µm dorsal length male, dg-4 in line with middle of dorsal muscle scars; China |
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– | Body >780 µm dorsal length female and >670 µm dorsal length male, dg-4 anterior to dorsal muscle scars |
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11 | Body elliptical, colorless to peach, small cheliceral fang (<33 µm) |
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– | Body rounded, grey to colorless, large cheliceral fang (>40 µm) |
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12 | Anterio-medial platelet wide (>140 µm) and more than or nearly twice as wide as long |
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– | Anterio-medial platelet unmodified (<140 µm) and less than twice as wide as long | ( |
13 | Anterio-medial and anterio-lateral platelets with consistent coloration (either colorless or colorled across) | ( |
– | Anterio-lateral platelets with coloration and anterio-medial platelet colorless |
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14 | Body in entirety conspicuously violet; currently known only from Arkansas |
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– | Body orange, red, pink, blue, or violet but not covering the majority of body |
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15 | Distribution east of the Great Plains, color violet to blue |
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– | Distribution within and west of the Great Plains |
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16 | Distribution in Washington or northern Oregon, orange to red |
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– | Distribution in and west of Great Plains excluding Washington and northern Oregon |
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17 | Violet to blue, females and males smaller than 600 and 500 µm, respectively | |
– | Collected outside of California, orange to red to clear, females and males larger than 600 and 500 µm, respectively | |
– | Collected in California and orange to clear | |
18 | Dorsal plate with large medial pores surrounded by distal ring of smaller pores (all pores uniform in other species), area posterior to coxal plate “bleached” in males |
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– | Dorsal plate with uniform pores, males without “bleached” area |
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19 | Anterio-lateral platelets with violet to blue coloration covering most of platelet |
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– | Anterio-lateral platelets with coloration restricted to the posterior half of platelet |
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20 | Females and males with dorsal lengths less than 575 and 450 µm, respectively |
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– | Females and males with dorsal lengths greater than 600 and 475 µm, respectively |
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Special thanks to Ian M. Smith (