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Species that have the parasitic larval stage suppressed illustrate that development of large eggs can obviate the need for larval feeding. However, the relative rarity of this phenomenon attests to the crucial role of larval dispersal in more derived water mite species.

Consistent with this view, many lentic water mites utilize far-flying hosts such as dragonflies e. However, if Torrenticola were utilizing such hosts, we would expect fewer instances of endemism, as well as more dispersal events across the east-west divide. Instead, all eastern Torrenticola diversity could be explained by as little as three dispersal events.

We speculate that this pattern can be explained by Torrenticola larval ecology. Torrenticola , like most lotic water mites, parasitize nematocerous flies, especially Chironomidae. In the proper habitat clean fast-flowing streams and riffles , Torrenticola adults are typically far more abundant than other water mites and larvae are easily identified by possessing fused coxal plates unlike all other Lebertioidea [Smith ] , which should increase both the likelihood of larvae being sampled on a host and being identified by researchers.

However, larval Torrenticola are rarely reported. Unfortunately, Torrenticola larvae are not currently correlated with adults, so there is no way to link species to hosts and we can only conclude that at least some unidentified species parasitize chironomids. This lack of knowledge has important ramifications because chironomids appear to have different inter- and intra-species natal fidelity. Although only a few studies have addressed adult chironomid dispersal e.

This trend is not restricted to chironomids as other aquatic insects that inhabit flowing water are also unable to disperse through dense riparian vegetation Titmus , Jackson and Resh , Peterson et al. Conversely, in areas with low vegetation, chironomids have been shown to disperse further Delettre and Morvan , and under such conditions can even cross into nearby catchments Krosch et al. We speculate that broadly distributed water mite species e. As discussed previously, loss of parasitism is especially likely with members of the Miniforma Group, which in addition to high endemism, also possess characters that have been proposed for species that have lost parasitism, such as smaller body size and a larger female genital opening Smith In either case host natal fidelity or loss of parasitism , the result is that species confined to a smaller geographic region may be more dependent on the adult stage for dispersal.

A sound understanding of Torrenticola larval ecology, including host associations, is critical to understanding the biology of the genus. Unfortunately, this area of inquiry is inhibited by our limited understanding, not only of water mites, but also chironomids, which are also understudied in North America, leaving most aspects of the biology of nearly all species involved a mystery.

To fully understand the evolution of Torrenticola within North America, analyses are needed that include worldwide taxon sampling and robust analyses of multiple genes, as well as a comprehensive understanding of larval ecology. To this end, we recommend the following areas of study for future investigation in North America: 1 surveys that correlate larval and adult chironomids; 2 dispersal studies on lotic chironomids; 3 surveys that correlate larval and adult water mites; and 4 surveys that investigate host specificity i.

Familial diagnosis. See Fisher et al. Subfamilial diagnosis. Type species. Generic diagnosis. Torrenticola anoplopalpa are similar to other members of the Rala Group T. Additionally, T. Torrenticola anoplopalpa sp. Dorsum — long; wide ellipsoid and colorless.

Anterio-medial platelets Anterio-lateral platelets long; Dgl-4 much closer to the edge of the dorsum than to the muscle scars distance between Dgl-4 Gnathosoma — Subcapitulum Rostrum long; Chelicerae long with curved fangs 40 long. Pedipalps short and stocky especially tibiae without extensions on femora and genua. Palpomeres: trochanter 35 long ; femur Venter — long; wide colorless. Gnathosomal bay Cxgl-4 subapical.

Medial suture 70 long. Genital plates long; wide. Additional measurements: Cx-1 long total ; long medial ; Cx-3 wide ; anterior venter long. Specific epithet anoplopalpa refers to the pedipalps, which lack tubercles on the femora and genua, an uncommon condition in Torrenticola , which usually have tuberculate ventral extensions anoplos , G.

Unfortunately, we were unable to acquire fresh material of Torrenticola anoplopalpa and so this species is not included in our phylogenetic analyses. However, we were able to examine material preserved in GAW for morphology. The overall appearance, incomplete hind coxal margins, lack of coloration, and distribution, are consistent with placing this species in the Rala Identification Group. Sterling Rd. Torrenticola arktonyx can be differentiated from all other Torrenticola by having distinctive longitudinal dark markings on the anterior portion of the dorsal plate that fade posteriorly.

Torrenticola arktonyx sp. Dorsum — — long; — wide ovoid with purple coloration separated into anterior and posterior portions with faint orange medially, also with distinctive longitudinal dark markings on the anterior portion of the dorsal plate that fade posteriorly. Anterio-lateral platelets — Dgl-4 much closer to the edge of the dorsum than to the muscle scars distance between Dgl-4 — Gnathosoma — Subcapitulum — Rostrum — long; 45— Chelicerae — long with curved fangs 45—55 45 long.

Pedipalps with dentate, flanged ventral extensions on femora and genua. Palpomeres: trochanter 40— Venter — — long; — wide with purple coloration. Medial suture 50—60 50 long. Genital plates — Additional measurements: Cx-1 — long total ; — long medial ; Cx-3 — wide ; anterior venter — Anterio-medial platelets — long; 55— Anterio-lateral platelets — long; 65—75 75 wide free from dorsal plate.

Gnathosoma — Subcapitulum — long ventral ; — long dorsal ; Rostrum — long; 35—40 40 wide. Chelicerae — long with curved fangs 40—45 40 long. Palpomeres: trochanter Gnathosomal bay — long; 65— Medial suture 75—90 80 long. Additional measurements: Cx-1 — long total ; — long medial ; Cx-3 — wide ; anterior venter — long. Unfortunately, we were unable to acquire fresh material of Torrenticola arktonyx and therefore this species is not included in our phylogenetic analyses.

We were able to examine material preserved in GAW for morphology, but due its unique characteristics, we are unable to place this species into either a species complex or identification group. However, based upon coloration, distribution, and gnathosomal shape, we speculate that future analyses will place this species in the Raptor Complex.

It is one of only four Eastern 2-Plates that have dark, bold, bluish-purple coloration also T. Torrenticola biscutella can be differentiated from T. Female T. Male T. Additionally, although T. Torrenticola biscutella sp. Dorsum — — long; — wide ovoid with bluish-purple coloration separated into anterior and posterior portions, and bold or faint orange medially. Anterio-lateral platelets — long; Rostrum — long; Chelicerae — long with curved fangs 55—70 56 long.

Pedipalps with tuberculate ventral extensions on femora and genua. Venter — — long; — wide with faint bluish-purple coloration. Medial suture absent. Genital plates Anterio-lateral platelets Dgl-4 much closer to the edge of the dorsum than to the muscle scars distance between Dgl-4 — Gnathosoma — Subcapitulum — long ventral ; — Rostrum 85— Chelicerae — long with curved fangs 45—50 46 long.

Gnathosomal bay — Medial suture 60—65 65 long. Additional measurements: Cx-1 — long total ; 99— long medial ; Cx-3 — wide ; anterior venter Specific epithet biscutella refers to the appearance of only two anterio-dorsal platelets due to the fusion of lateral platelets with the dorsal shield bi -, L. In all analyses, T.

The three of these species are morphologically similar to the more distantly-related T. Of these four species, the range of T. Based upon overall similarity, dorso-lateral platelet fusion, and distribution, we were able to place this species within the Eastern 2-Plate Identification Group.

Torrenticola bittikoferae are similar to other members of the Tricolor Group T. Torrenticola bittikoferae can be differentiated from most other Tricolor Complex except T. Torrenticola bittikoferae can be differentiated from T. Torrenticola bittikoferae male: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted. Dorsum — — long; — wide circular and colorless. Dgl-4 approaching midway between muscle scars and dorsum edge distance between Dgl-4 — Gnathosoma — Subcapitulum long ventral ; Rostrum 95— long; Chelicerae long with curved fangs 50 long short and conical.

Pedipalps with tuberculate ventral extensions with dentate tip on femora and tuberculate ventral extensions on genua. Venter — — long; — wide colorless. Medial suture Additional measurements: Cx-1 — long total ; Robert Crowell named the specific epithet bittikoferae after Lelia Bittikofer, his high school biology teacher. Torrenticola bittikoferae distribution. Unfortunately, we were unable to acquire fresh specimens of Torrenticola bittikoferae and therefore this species was not included in our phylogenetic analyses.

We were able to examine paratypes of two males, neither of which were dissected during slide preparation, making precise examination difficult. These specimens were remounted, but proper dissection risked fully damaging the specimen and was therefore avoided. The images in Figure 28A—B that appear to display a properly dissected specimen, were created by photographing each section dorsum and venter of the fully intact specimen and digitally editing the photographs so that the dorsum and venter could be easily compared with other species.

The overall appearance, short conical rostrum, and distribution of this species allows us to places it within the Tricolor Complex and Tricolor Identification Group. Torrenticola boettgeri K. Viets, a: Torrenticola esbelta Cramer, Torrenticola boettgeri are similar to other members of the Rala Group T. Note: measurements below are from the above three combined sources; those in parentheses are from the Guatemalan holotype Viets a as listed in Goldschmidt Torrenticola boettgeri female: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted.

Dorsum — — long; — wide ovoid, elongate, and colorless. Anterio-medial platelets — long; 38— Anterio-lateral platelets — long; 47—70 wide free from dorsal plate. Gnathosoma — Subcapitulum — long ventral ; long dorsal ; Rostrum 70— Chelicerae with curved fangs 65 long. Gnathosomal bay — long; 45—50 47 wide. Cxgl-4 apical. Medial suture 65—73 73 long. Genital plates — long; — wide. Additional measurements: Cx-1 long total ; — long medial ; Cx-3 wide ; anterior venter Torrenticola boettgeri male: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted.

Rostrum 80— Chelicerae — long with curved fangs Palpomeres: trochanter 30— Medial suture — long. Torrenticola boettgeri distribution. Blue star represents type locality Viets a ; blue diamond represents additional published record Cramer , as T. Unfortunately, we were unable to acquire fresh material of Torrenticola boettgeri and therefore this species is not included in our phylogenetic analyses. We were also unable to examine types, but were able to examine new material from New Mexico.

The overall appearance, incomplete hind coxal margins, distribution, and lack of coloration are consistent with placing this species in the Rala Identification Group. Viets a described T. Cramer described T. Cramer differentiated T. Furthermore, our material from New Mexico includes specimens with tibiae in between the previously recorded specimens Therefore, we consider T. They can be differentiated from Rusetria 4-Plates and T. Torrenticola bondi sp. Dorsum — — long; — wide ovoid with bluish-purple coloration separated into anterior and posterior portions and faint orange medially.

Dgl-4 approaching midway between muscle scars and dorsum edge distance between Dgl-4 — Gnathosoma — Subcapitulum — long ventral ; — long dorsal ; — tall faint bluish purple coloration. Rostrum — long; 50—55 Chelicerae — long with curved fangs 52—75 52 long. Gnathosomal bay — long; 90— Medial suture 10—15 10 long. Additional measurements: Cx-1 — long total ; — long medial ; Cx-3 — wide ; anterior venter Rostrum — long; 40—40 40 wide. Chelicerae — long with curved fangs 50—55 55 long.

Gnathosomal bay — Medial suture 55—70 60 long. Specific epithet bondi named in honor of arachnologist Jason Bond, for his research on species delimitation and integrative taxonomy, which has been an inspiration to JRF, and for his thoughtful career advice, which was greatly appreciated. Torrenticola bondi groups with other members of the Raptor Complex in all analyses with high support. However, the position of this clade varied with analysis.

Body proportions of male T. Torrenticola caerulea sp. Dorsum — — long; — wide ovoid with faint blue coloration anteriorly and posteriorly, broadly connected medially. Anterio-medial platelets — Chelicerae — long with curved fangs 62—70 65 long. Venter — — long; — wide with faint blue coloration. Gnathosomal bay — long; Additional measurements: Cx-1 Anterio-medial platelets 95— Rostrum Chelicerae — Additional measurements: Cx-1 — long total ; 85—95 95 long medial ; Cx-3 — wide ; anterior venter — long.

Specific epithet caerulea refers to the overall and diagnostic bluish appearance of this species caeruleus , L. The range of T. Torrenticola cardia are similar to other members of the Tricolor Complex T. Torrenticola cardia sp. Dorsum — — long; — wide ellipsoid with reddish-purple, bluish-purple or bright orange spot medially extending in a strip anteriorly often to the anterior-medial platelets. Anterio-medial platelets — long; 70—75 75 wide. Rostrum 90— long; 40— Chelicerae — long with curved fangs 60—60 60 long.

Pedipalps with stocky, tuberculate ventral extensions on femora and genua. Venter — — long; — wide with reddish-purple or bluish-purple coloration restricted to the edges of the gnathosomal bay, coxal plates, and genital plates. Medial suture 25—50 50 long.

Anterio-medial platelets — long; 65— Chelicerae Genital plates — long; Unfortunately, we were unable to acquire fresh material of Torrenticola cardia and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall appearance, short conical rostrum that is downturned in the male, and distribution, are consistent with placing this species in the Tricolor Complex and Tricolor Identification Group.

Torrenticola copipalpa are similar to members of the Miniforma group T. Torrenticola copipalpa sp. Dorsum — — long; — wide ovoid and usually colorless, occasionally with faint purple coloration without distinct pattern. Anterio-medial platelets — Chelicerae — long with curved fangs 50—59 55 long. Pedipalps short and stocky especially tibiae with broad, dentate, and anteriorly-directed ventral extensions on femora and dentate, flanged ventral extensions on genua. Palpomeres: trochanter 30—35 35 long ; femur 90— Medial suture 40—45 45 long.

Rostrum — long; 40— Chelicerae — long with curved fangs 45—65 55 long. Specific epithet copipalpa refers to the blade-like pedipalp femoral tubercles copis , L. This species overlaps with T. Based upon overall similarity, the pedipalp genu extensions, and western distribution, we were able to place this species in the Miniforma Identification Group. Torrenticola daemon are similar to other members of the Raptor Group T. Torrenticola daemon sp. Dorsum — — long; — wide circular with faint reddish-purple coloration separated into anterior and posterior portions, with bright reddish-purple coloration on the anterior-medial platelets, occasionally extending onto the dorsal plate.

Dgl-4 approximately halfway between the edge of the dorsum and the muscle scars distance between Dgl-4 — Gnathosoma — Subcapitulum — long ventral ; — long dorsal ; — Rostrum — long; 50—55 55 wide. Chelicerae — long with curved fangs 65—75 75 long.

Palpomeres: trochanter 50— Medial suture 10—15 15 long. Dorsum — — long; — wide circular with faint reddish-purple coloration separated into anterior and posterior portions. Gnathosoma — Subcapitulum — long ventral ; Chelicerae — long with curved fangs 60—65 60 long. Gnathosomal bay — long; 70—80 75 wide. Medial suture 50—55 55 long. Specific epithet daemon refers to the diagnostic red coloration on the anterio-medial platelets, which resemble the red eyes of an evil demon daemon , L.

Unfortunately, we were unable to acquire fresh material of Torrenticola daemon and therefore this species is not included in our phylogenetic analyses. The overall appearance, elongate subcapitular rostra, elongate pedipalpal tibiae, and Dgl-4 close to the muscle scars, are consistent with placing this species in the Raptor Complex and Raptor Identification Group.

Torrenticola danielleae are similar to other members of the Raptor Group T. Torrenticola danielleae sp. Dorsum — — long; — wide circular with reddish-purple coloration posteriorly extending in a strip anteriorly to the edge of the dorsal plate. Anterio-lateral platelets — long; 70—80 80 wide free from dorsal plate.

Gnathosoma — Subcapitulum — long ventral ; — long dorsal ; — tall colorless. Rostrum — long; 40—45 Chelicerae — long with curved fangs 50— Pedipalps with long, tuberculate ventral extensions on femora and genua. Palpomeres: trochanter 40—45 Venter — — long; — wide with reddish-purple coloration, occasionally faint.

Gnathosomal bay — long; 75—85 80 wide. Medial suture 25— Anterio-medial platelets — long; 45— Rostrum — long; 35—35 35 wide. Chelicerae — long with curved fangs 40—50 50 long. Medial suture 60—65 60 long. Specific epithet danielleae named in honor of Danielle Fisher—lab technician, environmental educator, colleague, friend, wife of JRF , and mother of Ivy, our beautiful daughter—for her tireless and immense contributions to this research, and for bettering the lives of all those around her.

Thank you, Danielle. Unfortunately, we were unable to acquire fresh material of Torrenticola danielleae and therefore this species is not included in our phylogenetic analyses. The overall appearance, elongate subcapitular rostra, and elongate pedipalpal tibiae, are consistent with placing this species in the Raptor Complex and the Raptor Identification Group.

Type series. Additionally, male T. Torrenticola delicatexa female: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted. Dorsum — — long; — wide ovoid with highly variable coloration, reddish-purple to purple occasionally bluish-purple separated into anterior and posterior portions.

Rostrum 90— long; Chelicerae long with curved fangs 45— Venter — — long; — wide usually colorless; occasionally with faint reddish-purple coloration. Medial suture 55—75 Genital plates — Additional measurements: Cx-1 — long total ; 78— long medial ; Cx-3 — Torrenticola delicatexa male: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted.

Anterio-medial platelets — long; Rostrum — long; 45—55 45 wide. Chelicerae — long with curved fangs 54—75 70 long. Additional measurements: Cx-1 — long total ; 93— long medial ; Cx-3 — wide ; anterior venter — long.

Habeeb did not offer an explanation for the specific epithet delicatexa and we are unable to offer helpful speculation. Torrenticola delicatexa distribution. We refrain from speculating on this COI variation, but consider this specimen as an outlier, and thus within our hypothesis for T. The position of this species was only strongly supported in our combined analysis, where it is recovered as sister to two other species: T.

However, T. Torrenticola dentirostra are similar to other members of the Nigroalba Group T. Torrenticola dentirostra sp. Dorsum — — long; — wide ovoid with faint purple coloration restricted posteriorly. Dgl-4 closer to the edge of the dorsum than to the muscle scars distance between Dgl-4 — Rostrum — long; 35—40 35 wide with dentate bump midway on dorsal edge.

Pedipalps with tuberculate ventral extensions dentate apically on femora and genua. Palpomeres: trochanter 30—35 30 long ; femur Venter — — long; — wide mostly colorless with faint purple genital plates. Gnathosomal bay 85— 85 long; 60—80 60 wide. Cxgl-4 far from apex and ventral. Medial suture 60—70 60 long.

Anterio-medial platelets — long; 40— Chelicerae — long with curved fangs 40—45 45 long. Medial suture 75—85 85 long. Additional measurements: Cx-1 — long total ; — Specific epithet dentirostra refers to the diagnostic tooth-like serrations on the dorsal surface of the rostrum dentis , L.

Unfortunately, we were unable to acquire fresh material of Torrenticola dentirostra and therefore this species is not included in our phylogenetic analyses. The overall similarity, small size, elongate subcapitular rostra, elongate pedipalpal tibiae, and dorsal purple coloration restricted to posterior half, place this species in the Raptor Complex and the Nigroalba Identification Group. Torrenticola dimorpha are similar to other members of the Tricolor Complex T.

Torrenticola dimorpha sp. Dorsum — — long; — wide colorless and ovoid. Anterior dorsal plate with medial extension covering nearly half the length of the anterio-medial platelets. Dgl-4 approximately halfway in between the edge of the dorsum and the muscle scars distance between Dgl-4 — Posterior dorsal apodeme long.

Rostrum 65—70 70 long; Chelicerae — long with curved fangs 60—70 70 long. Pedipalps with long, tuberculate ventral extensions on femora and no ventral extensions on genua. Medial suture 25—30 Muscle scars absent or very faint. Rostrum 45—50 Chelicerae — long with curved fangs 55—65 60 long. Pedipalps highly modified and expanded, with long, tuberculate ventral extensions on femora and no ventral extensions on genua.

Venter — — long; — wide colorless and highly modified with coxae II—IV forming a large ventral plate that covers the insertions of legs IV. Apodemes expanded internally. Genital plates — long; — wide triangular. Specific epithet dimorpha refers to the sexual dimorphism in terms of size and morphology di , G. In the combined analysis, T. This clade T.

This species is so unique that upon initial observation, specimens appear to be members of a different genus, especially the males. However, the short conical rostrum that is downturned in males is characteristic of other members of the Tricolor Identification Group. This species hypothesis is supported by biogeography, high divergence between species, and by highly distinctive morphological characteristics outlined in the diagnosis. Torrenticola dolichodactyla are similar to other members of the Rala Group T.

Torrenticola dolichodactyla sp. Dorsum — long; wide circular and colorless. Dgl-4 halfway between the muscle scars and the edge of the dorsum distance between Dgl-4 Gnathosoma — Subcapitulum long ventral ; long dorsal ; tall colorless. Rostrum long; 50 wide. Chelicerae long with curved fangs 65 long.

Pedipalps with elongate tarsi and tuberculate ventral extensions on femora and genua. Palpomeres: trochanter 55 long ; femur long ; genu 80 long ; tibia long; Gnathosomal bay long; wide. Medial suture 95 long. Specific epithet dolichodactyla refers to elongate pedipalp tarsus dolichos , G.

Unfortunately, we were unable to acquire fresh material of Torrenticola dolichodactyla and therefore this species is not included in our phylogenetic analyses. The overall appearance, incomplete hind coxal margins, distribution, and lack of color are consistent with placing this species in the Rala Identification Group. Torrenticola dunni sp. Dorsum — — long; — wide ovoid with purple or reddish-purple coloration separated into anterior and posterior portions, and occasionally with faint orange medially.

Rostrum — long; 45—50 45 wide. Venter — — long; — wide with faint bluish-purple or reddish purple coloration or colorless. Medial suture 20—25 20 long. Rostrum — Chelicerae — long with curved fangs 50—55 Gnathosomal bay — long; 75— Specific epithet dunni named in honor of Rob Dunn of North Carolina State University, for his exceptional writings and research that personalize ecology by bringing nature indoors; and particularly for his storytelling ability, in which he wonderfully conveys that we humans, rather than being separate from nature, are indeed just as wild as what we perceive outdoors—a sentiment exemplified by his book, Wild Life of Our Bodies: Predators , Parasites , and Partners That Shape Who We Are Today Given our collection efforts across the Appalachians, it is reasonable to speculate that T.

Atractides ellipsoidalis : Marshall : Torrenticola rectiforma Habeeb, 1. Torrenticola ellipsoidalis are similar to other members of the Ellipsoidalis Group T. Torrenticola ellipsoidalis female: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted. Dorsum — — long; — wide rectangular and usually colorless, occasionally with faint purple coloration without distinct pattern.

Chelicerae — long with curved fangs 61—74 long. Medial suture 40— Additional measurements: Cx-1 — Torrenticola ellipsoidalis male: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted. Anterio-lateral platelets — long; 85— Chelicerae — long with curved fangs 60—74 long.

Palpomeres: trochanter 35— Medial suture 70—90 long. We expand the range into most of western North America. Torrenticola ellipsoidalis distribution. Red dots indicate material examined. This is higher sequence variability than in many species hypotheses presented herein.

An exception is that a single specimen DNA is 2. This specimen is indistinguishable from other specimens and is collected from the same location. We do not find evidence to propose it as a separate species and therefore include it within T. Based upon overall similarity, body size, and distribution, we place this species within the Ellipsoidalis Identification Group.

Upon examining the types of T. Furthermore, the main character Habeeb used to differentiate T. Therefore, we synonymize T. Additionally, they can be differentiated from Rusetria 4-Plates and T. Dorsum — — long; — wide ovoid and elongate with purple coloration separated into anterior and posterior portions.

Pedipalps with tuberculate ventral extensions on femora and on genua. Gnathosoma — Subcapitulum — long ventral ; — long dorsal ; 85— Chelicerae — long with curved fangs 34—42 42 long. Gnathosomal bay — long; 50—55 55 wide. Medial suture 55—80 55 long. Additional measurements: Cx-1 — long total ; 82—98 98 long medial ; Cx-3 — wide ; anterior venter — long. Specific epithet elongata refers to elongated bodies of this species, which is more pronounced than in all other North American Torrenticola elongatus , L.

Based upon overall similarity, an elongate body, and distribution, we place this species in the Elongata Identification Group. Torrenticola elusiva are similar to other members of the Raptor Group T. Additionally, two character systems that vary minimally between sexes are rostrum and pedipalp tibiae proportions, which do differ between T. Dorsum — long; wide circular with bluish-purple coloration posteriorly with a broad anterior extension reaching the anterior edge of the dorsal plate.

Dgl-4 much closer to the muscle scars than to edge of dorsum distance between Dgl-4 Gnathosoma — Subcapitulum long ventral ; long dorsal ; Chelicerae long with curved fangs 59 long. Pedipalps elongate especially tibiae with tuberculate ventral extensions with dentate tip on femora and tuberculate ventral extensions on genua.

Additional measurements: Cx-1 long total ; long medial ; Cx-3 wide ; anterior venter Specific epithet elusiva refers to the fact that we were only able to find a single specimen of this species, despite extensive searching among the abundant samples taken from the type locality in New Brunswick.

Torrenticola elusiva groups with other members of the Raptor Complex in all analyses with high support. Furthermore, this species is known from only a single female, so morphological variation could not be investigated. However, this specimen was different enough in terms of morphology and sequence to warrant a separate description. We place this species within the Raptor Identification group based upon similarity with those species.

Torrenticola erectirostra are similar to other members of the Erectirostra Group T. They can be differentiated from all other Torrenticola , except T. Torrenticola erectirostra sp. Dorsum — — long; — wide ovoid with bluish-purple or purple coloration separated into anterior and posterior portions with orange medially.

Rostrum — long; 55— Pedipalps short and stocky especially tibiae with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter 45—55 50 long ; femur — Gnathosomal bay — long; — wide. Cxgl-4 far from apex. Rostrum 90— Chelicerae — long with curved fangs 45—50 50 long. Medial suture 75— Specific epithet erectirostra refers to the upturned rostrum characteristic of members of the Erectirostra Group erectus , raised up; rostrum , L.

Torrenticola erectirostra groups with other members of the Raptor Complex with high support and specimens are 2. This variation in COI is higher than in most species hypotheses proposed herein, especially since those specimens were form the same region New Brunswick. However, we could not find morphological differences that corresponded to clades in our analysis, and we were only able to examine four sequence, therefore, we consider these specimens to represent a single species hypothesis.

This species groups with T. This species hypothesis is supported by biogeography, molecular divergence although COI variation is greater than most hypotheses herein , and by the morphological characters outlined in the diagnosis. Torrenticola feminellai sp. Dorsum — — long; — wide circular with reddish-purple coloration in the shape of an hourglass.

Chelicerae — long with curved fangs 65—75 70 long. Anterio-lateral platelets — long; 55— Chelicerae — long with curved fangs 50—60 55 long. Palpomeres: trochanter 35—45 Venter — — long; — colorless. Medial suture 65—75 70 long.

Specific epithet feminellai named in honor of Jack Feminella, professor of biology at Auburn University, who believed in me JRF enough to employ me as a lab technician in stream ecology, write a winning recommendation letter for graduate school, and was the first to teach me how to conduct self-directed research.

Unfortunately, we were unable to acquire fresh material of Torrenticola feminellai and therefore this species is not included in our phylogenetic analyses. The overall similarity, distribution in the east, and fusion of the dorso-lateral platelets to the dorsal plate, are consistent with placing this species in the Rusetria Complex and the Eastern 2-Plate Identification Group. Torrenticola flangipalpa are similar to other members of the Nigroalba Group T.

Other Torrenticola with purple dorsal coloration restricted posteriorly, such as T. Torrenticola flangipalpa sp. Dorsum — — long; — wide ovoid with purple or bluish-purple coloration restricted posteriorly. Gnathosoma — Subcapitulum — Chelicerae — long with curved fangs 43—50 44 long. Pedipalps elongate especially tibiae with broad, dentate, anteriorly-directed flange on femora and with variable, dentate flange-like extension on genua.

Venter — — long; — wide with faint purple or bluish-purple coloration. Chelicerae — long with curved fangs 39—62 40 long. Specific epithet flangipalpa refers the expanded i. That clade of three species corresponds to an identification group, Nigroalba Group, the members of which are easily differentiated by their size, coloration, long medial suture in females, and overall appearance. Torrenticola folkertsae are similar to other members of the Partial 2-Plate Group T.

Torrenticola folkertsae sp. Dorsum — — long; — wide ovoid with faint reddish-purple medially. Anterio-lateral platelets — long; 85—95 85 wide partially fused to dorsal plate. Chelicerae — long with curved fangs 65—70 70 long. Medial suture 15—20 20 long.

Anterio-medial platelets — long; 55—65 55 wide. Chelicerae — long with curved fangs 55— Additional measurements: Cx-1 — long total ; — long medial ; Cx-3 — wide ; anterior venter — Specific epithet folkertsae named in honor of Debbie Folkerts, professor of biology at Auburn University, who, together with her late husband George Folkerts, were instrumental to JRF in channeling his passion for natural history and teaching into a career path. Unfortunately, we were unable to acquire fresh material of Torrenticola folkertsae and therefore this species is not included in our phylogenetic analyses.

The overall similarity, distribution, and partial fusion of the dorso-lateral platelets to the dorsal plate, are consistent with placing this species in the Rusetria Complex and the Partial 2-Plate Identification Group. Torrenticola glomerabilis sp.

Dorsum — — long; — wide circular with bold bluish-purple or reddish-purple coloration separated into anterior and posterior portions. Chelicerae — long with curved fangs 50—55 53 long. Venter — — long; — wide bold bluish-purple or reddish-purple coloration. Rostrum — long; 35—40 35 wide. Palpomeres: trochanter 35—35 35 long ; femur 95— Medial suture 85— Additional measurements: Cx-1 — long total ; 88— long medial ; Cx-3 — wide ; anterior venter — long.

Specific epithet glomerabilis refers to the rounded body of this species compared to all other members of the Rusetria Complex glomerabilis , L. Of these species, T. Because of this, we place T. Their unique shape and interesting phylogenetic affinity flag this species as important to future studies on the evolution of eastern members of the Rusetria Complex.

Torrenticola gnoma are similar to other members of the Raptor Group T. Additionally, female T. Torrenticola gnoma sp. Dorsum — — long; — wide circular with a large spot of coloration medially extending in a thin strip anteriorly to the edge of the dorsal plate, coloration variable from navy blue to purple to pink. Dgl-4 much closer to the muscle scar than to dorsum edge distance between Dgl-4 — Chelicerae — long with curved fangs 53—60 54 long.

Pedipalps elongate with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter 40—45 40 long ; femur — Medial suture 15— Additional measurements: Cx-1 — long total ; 84— long medial ; Cx-3 — wide ; anterior venter — Anterio-medial platelets 1.

Gnathosoma— Subcapitulum — long ventral ; — long dorsal ; Chelicerae — long with curved fangs 41—51 50 long. Additional measurements: Cx-1 — long total ; 90— long medial ; Cx-3 — wide ; anterior venter Specific epithet gnoma refers to the dorsal pattern, which, although variable, resembles the head and cap of a gnome gnoma , L.

Based upon overall similarity, distribution, and phylogenetic position, this species is placed within the Raptor Identification Group. Torrenticola gorti specimens from Tellico River system in Monroe County Tennessee can be differentiated from all other Torrenticola by the distinctively dark coloration with a red spot dorsally. Torrenticola gorti sp.

Dorsum — — long; — wide ovoid and elongate with three distinct color morphs: 1 navy blue coloration separated into anterior and posterior portions that meet or nearly meet laterally, and with bold orange coloration in between, from Tellico River, Monroe County, Tennessee; 2 purple coloration separated into anterior and posterior portions; 3 purple coloration separated into anterior and posterior portions with a strip of bold orange medially.

Chelicerae — long with curved fangs 56—60 60 long. Dorsum — — long; — wide ovoid and elongate with three distinct color morphs: 1 navy blue coloration separated into anterior and posterior portions that meet or nearly meet laterally, and with bold orange coloration in between; 2 purple coloration separated into anterior and posterior portions; 3 purple coloration separated into anterior and posterior portions with a strip of bold orange medially.

Chelicerae — long with curved fangs 36—50 50 long. Venter — — long; — wide with three distinct color morphs: 1 navy-blue coloration; 2 colorless; 3 purple coloration. In the film, Gort was depicted with a dark body and a single red eye that shot a destructive beam. Members of this species can be highly variable in color. Some members have reddish-purple or purple dorsal coloration that is separated into anterior and posterior portions.

Ventral coloration can be bold, faint, or absent. Members from Tellico River, Monroe County Tennessee , can be readily differentiated from all other Torrenticola by being dark navy blue with a red dorsal oval. Torrenticola hoosieri are similar to other members of the Tricolor Complex T. Torrenticola hoosieri sp. Dorsum — — long; — wide ellipsoid and colorless, occasionally with pink coloration without a distinct pattern.

Anterio-medial platelets — long; 55—60 55 wide. Chelicerae — long with curved fangs 75—77 75 long. Pedipalps without extensions on femora and genua. Palpomeres: trochanter 50—55 50 long ; femur — Medial suture 30—30 30 long. Dorsum — — long; — wide ellipsoid and colorless. Chelicerae — long with curved fangs 65—70 Gnathosomal bay — long; 80—85 85 wide.

Specific epithet hoosieri refers to Hoosier, the English demonym for a person from Indiana, the type locality. This species is placed in the Tricolor Identification Group. Atractides indistinctus Marshall, Torrenticola indistincta female: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted.

Dorsum — — long; — wide ovoid with faint coloration separated into anterior and posterior portions and connected medially. Chelicerae — long with curved fangs 60—80 Palpomeres: trochanter 45—65 Torrenticola indistincta male: A dorsal plates B venter legs removed C subcapitulum D pedipalp setae not accurately depicted. Rostrum 85— 85 long; Medial suture 65—75 75 long. Young reported this species from Colorado, but this likely represents T.

Torrenticola indistincta distribution. Blue diamonds indicate previously published reports. Unfortunately, we were unable to acquire fresh material of Torrenticola indistincta and therefore this species was not included in our phylogenetic analyses.

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