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. 2016 Apr 21:(582):13-110.
doi: 10.3897/zookeys.582.7684. eCollection 2016.

Systematics of testudacarine torrent mites (Acari, Hydrachnidia, Torrenticolidae) with descriptions of 13 new species from North America

Joseph C O'Neill  1 , J Ray Fisher  1 , Whitney A Nelson  1 , Micheal J Skvarla  1 , Danielle M Fisher  1 , Ashley P G Dowling  1
Affiliations

Systematics of testudacarine torrent mites (Acari, Hydrachnidia, Torrenticolidae) with descriptions of 13 new species from North America

Joseph C O'Neill et al. Zookeys. .

Abstract

Thirteen new species of North American Testudacarus (Torrenticolidae: Testudacarinae) are described: Testudacarus deceptivus O'Neill & Dowling, sp. n., Testudacarus hitchensi O'Neill & Dowling, sp. n., Testudacarus harrisi O'Neill & Dowling, sp. n., Testudacarus dennetti O'Neill & Dowling, sp. n., Testudacarus dawkinsi O'Neill & Dowling, sp. n., Testudacarus radwellae O'Neill & Dowling, sp. n., Testudacarus kirkwoodae O'Neill & Dowling, sp. n., Testudacarus hyporhynchus O'Neill & Dowling, sp. n., Testudacarus smithi O'Neill & Dowling, sp. n., Testudacarus rollerae O'Neill & Dowling, sp. n., Testudacarus elongatus O'Neill & Dowling, sp. n., Testudacarus rectangulatus O'Neill & Dowling, sp. n., and Testudacarus oblongatus O'Neill & Dowling, sp. n. Testudacarus vulgaris Habeeb, 1954 is resurrected from synonymy with Testudacarus minimus and redescribed. Debsacarus (Habeeb, 1961), Testudacarus americanus Marshall, 1943, and Testudacarus minimus Marshall, 1943 are redescribed. All redescriptions are from original types. Species delimination was accomplished through examination of morphology, biogeography, and molecular phylogenetics of the barcoding region of COI. Other species are addressed and a key to world species is presented. For Testudacarinae, this represents the first published: 1) descriptions from multiple specimens (i.e. intraspecific variation); 2) colored photographs; 3) explicit illustrations and discussion of sexual dimorphism within the subfamily; 4) genetic data. A comprehensive testudacarine reference list is also included.

Keywords: Debsacarus; Hydrachnidia; Hydrachnidiae; Testudacarinae; Testudacarus; water mites.

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Figures

Figure 1.
Figure 1.
Testudacarine male dorsum (generalized): (Left) – anterio-medial platelet (amp); anterio-lateral platelet (alp); dorsal plate (dp); lateral platelets (lp); (Right) – dorso-glandularia (dg); post-ocularial setae (pos); dorsal membrane (dm); lyriffisures (l); muscle scars (ms); latero-glandularia (lg).
Figure 2.
Figure 2.
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).
Figure 3.
Figure 3.
Testudacarine male venter (generalized): Left – coxo-glandularia (cg); latero-glandularia (lg); ventro-glandularia (vg); Middle – coxae (c). Right – gnathosomal bay (gb); coxae-II+III midline (ml); genital field (gf); acetabula (a); line of secondary sclerotization (ss); excretory pore (ep).
Figure 4.
Figure 4.
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).
Figure 5.
Figure 5.
Testudacarine gnathosoma (generalized): chelicerae (c); posterio-dorsal apodeme (pda); posterio-ventral apodeme (pva); subcapitulum (sc); fang (f); rostrum (r).
Figure 6.
Figure 6.
Testudacarinae molecular phylogeny and species complexes: (Left) combined 28S and COI Bayesian analysis resulting in a five branched soft polytomy (●: >95% posterior probability); monophyly tested across Torrenticolidae but not depicted; (A–E represent tree continuation in Figs 8, 12, 23, 32, and 43 respectively; (Right) species complexes with illustrative descriptions.
Figure 7.
Figure 7.
Testudacarine sexual dimorphism: female dorsal shield (A) and ventral shield (C) differing from male (B, D) by the following characters: 1) dorso-glandularia-4 positioned far closer to muscle scares; 2) area of secondary sclerotization always present (males rarely present; very small if present); 3) with shorter coxae-II+III midline; 4) genital field enveloped by coxal field; 5) larger and rounder body (males around 80% of female size); 6) excretory pore well separated from ventral line of secondary sclerotization.
Figure 8.
Figure 8.
Debsacarus oribatoides molecular phylogeny: 28S and COI Bayesian analysis showing strong support single distinct clade (●: >95% posterior probability); clade exhibits <.6% divergence in COI within and >15% divergence between any other clade (not pictured); continuation of (E) lineage from Fig. 6.
Figure 9.
Figure 9.
Debsacarus oribatoides gnathosoma (generalized).
Figure 10.
Figure 10.
Debsacarus oribatoides female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 11.
Figure 11.
Debsacarus oribatoides male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 12.
Figure 12.
Testudacarus minimus complex molecular phylogeny: 28S and COI Bayesian analysis showing strong support for a soft polytomy with three distinct clades (●: >95% posterior probability); colored clades exhibit <2.5% divergence in COI within and >6.5% divergence between; continuation of (A) lineage from Fig. 6.
Figure 13.
Figure 13.
Testudacarus minimus female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 14.
Figure 14.
Testudacarus minimus color variation.
Figure 15.
Figure 15.
Testudacarus minimus male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 16.
Figure 16.
Testudacarus vulgaris female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 17.
Figure 17.
Testudacarus vulgaris color variation.
Figure 18.
Figure 18.
Testudacarus vulgaris male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 19.
Figure 19.
Testudacarus deceptivus female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 20.
Figure 20.
Testudacarus deceptivus male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 21.
Figure 21.
Testudacarus radwellae female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 22.
Figure 22.
Testudacarus radwellae male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 23.
Figure 23.
Testudacarus hitchensi complex molecular phylogeny: 28S and COI Bayesian analysis showing strong support for at least four distinct clades, but suggesting more (●: >95% posterior probability); excepting green clade, clades exhibit <1.5% divergence in COI within and >6% between; green clade exhibits <4.5% within and >9.5% between other clades; specimens in red constitute additional suspected species based on genetic data, but lack morphological or distributional variation from green clade; continuation of (B) lineage from Fig. 6.
Figure 24.
Figure 24.
Testudacarus hitchensi female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 25.
Figure 25.
Testudacarus hitchensi male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 26.
Figure 26.
Testudacarus harrisi female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 27.
Figure 27.
Testudacarus harrisi male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 28.
Figure 28.
Testudacarus dennetti female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 29.
Figure 29.
Testudacarus dennetti male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 30.
Figure 30.
Testudacarus dawkinsi female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 31.
Figure 31.
Testudacarus dawkinsi male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 32.
Figure 32.
Testudacarus americanus complex molecular phylogeny: 28S and COI Bayesian analysis showing strong support for five distinct clades (●: >95% posterior probability); excluding pink clade, colored clades exhibit <1.3% divergence in COI within and >9% divergence between; pink exhibits 4.5% variation within; red specimen is a suspected species based on genetic data, but specimen is teneral and too badly damaged to diagnose; continuation of (C) lineage from Fig. 6.
Figure 33.
Figure 33.
Testudacarus americanus female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 34.
Figure 34.
Testudacarus americanus male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 35.
Figure 35.
Testudacarus kirkwoodae female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 36.
Figure 36.
Testudacarus hyporhynchus sp. n. gnathosoma (generalized).
Figure 37.
Figure 37.
Testudacarus hyporhynchus female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 38.
Figure 38.
Testudacarus hyporhynchus male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 39.
Figure 39.
Testudacarus smithi female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 40.
Figure 40.
Testudacarus smithi male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 41.
Figure 41.
Testudacarus rollerae female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 42.
Figure 42.
Testudacarus rollerae male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 43.
Figure 43.
Testudacarus elongatus complex molecular phylogeny: 28S and COI Bayesian analysis showing strong support at least three distinct clades (●: >95% posterior probability); colored clades exhibit <2.4% divergence in COI within and >3.3% divergence between; divergence of the two basal clades >9%; continuation of (D) lineage from Fig. 43.
Figure 44.
Figure 44.
Testudacarus elongatus female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 45.
Figure 45.
Testudacarus elongatus male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 46.
Figure 46.
Testudacarus rectangulatus male: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 47.
Figure 47.
Testudacarus oblongatus female: (Left) dorsum; (Right) venter. Scale: 100 µm.
Figure 48.
Figure 48.
Testudacarus oblongatus male: (Left) dorsum; (Right) venter. Scale: 100 µm.
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