Submitted: April 23th, 2020 – Accepted: June 26th, 2020 – Published online: June 28th, 2020
To link and cite this article:
doi: 10.5710/AMGH.26.06.2020.3354
1 MITES (ACARI, ORIBATIDA, NANHERMANNIDAE) FROM THE EOCENE
2 OF PATAGONIA: FIRST SOUTHERN HEMISPHERE FOSSIL RECORD IN
3 MARINE SEDIMENTS
4 ÁCAROS (ACARI, ORIBATIDA, NANHERMANNIDAE) DEL EOCENO DE
5 PATAGONIA: PRIMER REGISTRO FÓSIL DEL HEMISFERIO SUR EN
6 SEDIMENTOS MARINOS
7
8 DAMIÁN A. FERNÁNDEZ1, PABLO A. MARTÍNEZ2, LUIS PALAZZESI3,
9 VIVIANA D. BARREDA3.
10 1 Laboratorio de Geomorfología y Cuaternario, CADIC-CONICET, Bernardo Houssay
11 200, V9410 Ushuaia, Argentina. [email protected]
12 2 Departamento de Biología, Facultad de Ciencias Exactas y Naturales, Universidad
13 Nacional de Mar del Plata, Deán Funes 3350, B7602AYL Mar del Plata, Argentina.
15 3 Sección Paleopalinología, División Paleobotánica, MACN-CONICET, Av. Ángel
16 Gallardo 470, C1405DJR Buenos Aires, Argentina. [email protected];
18
19 14 pag. (text + references); 2 figs.
20
21 FERNÁNDEZ ET AL.: FOSSIL MITES FROM SW PATAGONIA
22 First record of a pre-Pleistocene fossil mite (Acari, Oribatida, Nanhermannidae) from
23 the Southern Hemisphere in marine sediments.
24
25 Corresponding author: Damián A. Fernández, [email protected] 1
26 Key words. Mites. Oribatida. Middle/Late Eocene.. Patagonia. Argentina.
27 Palabras clave. Ácaros. Oribatida. Eoceno Medio/Superior. Patagonia. Argentina.
28
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29 COSMOPOLITAN, ABUNDANT, AND ADAPTED to nearly all terrestrial environments
30 today, mites are, however, uncommon in the fossil record. Extant diversity of mites
31 is second only to that of insects. With some 50,000 species described, of which
32 nearly 25% are oribatids, their past diversity is not well reflected in the fossil record
33 because of their rarity as fossils, especially in marine deposits and older strata
34 (Selden et al., 2008). Mites are more abundant in Quaternaries sites (e.g. Woolley,
35 1969; Elias, 1994; Heyne & Coetzee, 2001) and are being used in biostratigraphic
36 and palaeoclimatic studies of that period (e.g. Erickson, 1988; Schelvis, 1990;
37 Erickson et al., 2003; Krivolutsky & Sidorchuk, 2003; Mauquoy & van Geel, 2007;
38 Demske et al., 2013). A few are known from Cenozoic ambers (e.g. Selden et al.,
39 2008 and references there in; Dunlop et al., 2013; Khaustov, 2014; Sidorchuk et al.,
40 2019; Klimov et al., 2019; Poinar, 2019; Stilwell et al., 2020). Mites from
41 Cretaceous ambers are far fewer in number (e.g., Selden et al., 2008 and references
42 there in; Judson & Mąąkol, 2009; Sidorchuk et al., 2015b; Sidorchuk & Behan-
43 Pelletier, 2017; Arillo et al., 2018 and references there in). The only record from
44 South America until now is a large parasitengonid mite (Acari, Erythraeoidea) from
45 the continental Early Cretaceous Crato Formation of Brazil (Dunlop, 2007). There
46 are known three oribatids from the Jurassic (Krivolutsky & Krassilov, 1977; Sivhed
47 & Wallwork 1978; Selden et al., 2008), and only two species from Triassic amber,
48 north-eastern Italy (Sidorchuk et al., 2015a). The oldest known mite fossils come
49 from the Devonian localities of Gilboa, USA (Norton et al., 1988; Kethley et al.,
50 1989) and Rhynie, Scotland (Hirst, 1923).
51 In this note we report the finding of a fossil remain belonging to an oribatid mite. It
52 is a leg, morphologically assignable to the family Nanhermannidae. It is the first
53 record of a pre-Pleistocene fossil mite from the Southern Hemisphere in marine
3
54 sediments. The specimen comes from the Río Turbio Formation (RTF), SW of Santa
55 Cruz province, Patagonia, Argentina (Fig. 1). The RTF represents sedimentation in a
56 transitional environment during the Middle Eocene, period when a global climate
57 event called Middle Eocene Climatic Optimum (MECO) is recorded (González
58 Estebenet et al., 2015). This finding supports the presence of these mites in the
59 southernmost tip of South America during the Eocene. Also provides a new record,
60 important from the phylogenetic and paleoecological point of view.
61 Institutional abbreviations. CADIC, Centro Austral de Investigaciones Científicas,
62 Ushuaia, Argentina; CONICET, Consejo Nacional de Investigaciones Cinetíficas y
63 Técnicas, Buenos Aires, Argentina; MACN, Museo Argentino de Ciencias Naturales
64 “Bernardino Rivadavia”, Buenos Aires, Argentina.
65 Anatomical abbreviations. Fe, femur; Ge, genu; Ta, tarsus; Ti, tibia; Tr,
66 trochanter.
67 MATERIALS AND METHODS
68 The fossil specimen was recognized from the analysis of palynological slides
69 deposited in the Museo Regional Provincial Padre Manuel Jesús Molina (Río
70 Gallegos, Santa Cruz), under the acronym MPM-PB, and catalog number 21662. The
71 sample analyzed corresponds to a dark shale that comes from an outcrop in the
72 surroundings of Río Turbio city (Fig. 1) and belongs to the upper member of the RTF
73 (Romero, 1977). The techniques for palynological processing comes from Wood et
74 al. (1996). The specimen was studied using a Leica DM2500 light microscope and
75 photographed using a Leica DFC450 C camera. The coordinate of the specimen
76 corresponds to the England Finder ruler. The morphological description follows the
77 terminology of Norton & Behan-Pelletier (2009). The fossil was drawn to obtain a
78 morphological detail by a compound microscope Olympus CX 31 with drawing
4
79 tube. For fossil identification Fujikawa (1990) and Ermilov (2009) were consulted.
80 SYSTEMATIC PALEONTOLOGY
81 Class ARACHNIDA Lamark, 1801
82 Subclass ACARI Leach, 1817
83 Order SARCOPTIFORMES Reuter, 1909
84 Suborder ORIBATIDA van der Hammen, 1968
85 Family NANHERMANNIDAE Sellnick, 1928
86 Referred material. Palynological slide MPM-PB 21662, O47–3. Museo Regional
87 Provincial Padre Manuel Jesús Molina, Río Gallegos, Santa Cruz, Argentina.
88 Geographic occurrence. Surroundings of Río Turbio city, 5 km northeast of the city,
89 beside National Route 40, Santa Cruz Province, Patagonia, Argentina (51° 30' 37.00" S
90 72° 15' 33.00" W) (Fig. 1).
91 Stratigraphic occurrence. Río Turbio Formation (Middle/Late Eocene, Priabonian, ca.
92 40 Myr) (González Estebenet et al., 2015).
93 Description. Tarsus almost complete, part of the genu, and complete femur and
94 trochanter. Tibia, in this group similar in size to the genu, covered by organic remains.
95 Length of each segment: Ta = 43 µm, Ti + Ge = 28 µm (where Ti is equal to or slightly
96 greater than Ge), Fe = 33 µm, Tr = 18 µm. In the tarsus, claw and numerous phaneres
97 are observed, but common setae and solenidia are indistinguishable. Some setae overlap
98 with their pair. Dorsal setae united in groups of two or more. Anterior ventral seta
99 stands out (probably s seta), apparent “holes” observed inside, possible eupathidium
100 (fanera that, unlike a normal seta that has a dense core of chitin, has a channel along its
101 axis). Two dorsal phaneres distinguished in the visible part of the genu, which would
102 correspond to a seta and a solenidium, and a ventral one. Dorsal and a lateral seta seen
103 on the femur. Small circles, possible insertions of phaneres or taphonomic artifacts.
5
104 Ventrolateral seta in the trochanter (Fig. 2).
105 Remarks. The leg has been completely preserved. The different segments can be
106 distinguished. In some areas there are overlapped organic matter remains. While the
107 cuticle of the leg has been altered becoming almost transparent (probably due to the
108 palynological processing), the setae remain in their position and are brown. Tarsus setae
109 are the darkest and more rigid part. Towards the base of the leg, they appear lighter and
110 somewhat collapsed (Fig. 2).
111 DISCUSSION AND CONCLUSIONS
112 The morphology and arrangement of the setae allow us to assign the mite remain here
113 described to the Nanhermannidae family. Although the description of the legs is not
114 frequent in representatives of Nanhermannidae, widened setae like those of the leg
115 described here are typical of Nanhermannia species (e.g. Fujikawa, 1990 Figs. 3, 4;
116 Ermilov, 2009 Fig. 4). This finding confirms the presence of mites related to that family
117 in the southern tip of South America during the Middle/Late Eocene. The most closely
118 related living clades to the RTF specimen are currently distributed in South America,
119 mainly in Venezuela, Ecuador, Peru, Chile, Brazil, Bolivia, Uruguay and Argentina. In
120 Patagonia, extant species have been cited for El Bolsón (Balogh & Csiszar, 1963),
121 Punta Bandera (Hammer, 1962b), Grand Malvina Island (Starý & Block, 1996)
122 (Argentina), Puerto Montt (Hammer, 1962a ), Punta Arenas and Puerto Natales
123 (Ermilov, 2016) (Chile).
124 Two main hypotheses can be raised about the living environment of the fossil mite: 1)
125 mite lived in a littoral environment 2) mite lived in leaf litter.
126 Hypothesis 1 is supported by the fact that the fossil bearing sediments were deposited in
127 a shallow marine environment according to sedimentological (Rodríguez Raising, 2015)
128 and paleontological studies (González Estebenet et al., 2012). While extant family
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129 records mentioned above were made in terrestrial biotopes, Colloff (1983) cited the
130 presence of a species of Nanhermannia Berlese, 1913 in coastal lichens in the Islands of
131 the Clyde, Scotland. The leg shows signs of alteration as a result of the palynological
132 processing (see Remarks). The non-preserved mite body remains could have been lost
133 due to this process (and not due to transport). The leg is articulated and very well
134 preserved with the occurrence of delicate structures, hence we could assume that it has
135 not been transported far from the original habitat.
136 A continental origin of the fossil mite (Hypotheses 2) is supported by the fact that most
137 of the extant Nanhermannidae species live in leaf litter (Norton & Behan-Pelletier,
138 2009). In that case, the fossil leg may have been transported by water with the pollen
139 and spores also found in the sample.
140 In the RTF we also found an important number of claws (180 in 53 studied samples,
141 maximum peak =18) similar to that of the specimen here studied. According to
142 numerous studies on oribatid ecology (Erickson, 1988; Labandeira et al., 1997;
143 Erickson et al., 2003) they have an important potential as paleoenvironmental indicators
144 of energy of the depositional environment. Future works on these claws would help to
145 better understand their importance as paleoenvironmental indicators.
146 This finding and the associated claws assigned to the family Nanhermannidae in
147 paleopalynological slides adds an important taxon to the group of non-sporopollinic
148 palynomorphs.
149 ACKNOWLEDGMENTS
150 The authors wish to express their thanks to: Amalia González for the stratigraphic
151 column; Orlando Cárdenas for technical work; Pamela Alli for helping with samples;
152 Dr. Adolfina Savoretti for helping with writing; Dr. Roy Norton for his comments and
153 suggestions; and four anonymous reviewers who significantly improved the quality of
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154 our manuscript. Financial support was provided by Consejo Nacional de Investigaciones
155 Científicas y Técnicas (PIP 2014–0259) and Agencia Nacional de Investigaciones
156 Científicas y Técnicas (PICT 2017–0671).
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289 Figure captions
290 Figure. 1. Localization and geologic details of the sample studied.
291 Figure. 2. 1, Light microscope image; 2, Tube drawing of the specimen. Fa, fanera; Cl,
292 claw; Fe, femur; Ge, genu; Se, seta; Ti, tibia; Tr, trochanter; Ta, tarsus; (a), antelateral
293 setae; (u), unguinal setae; s, subunguinal seta; p, proral seta; (tc), tectal setae; (pv),
294 primiventral setae; v, ventral seta; l, lateral seta; d, dorsal seta; ω, tarsal solenidium; δ,
295 genual solenidium. Parentheses indicate setal pair. Scale bars equals 20 µm.
296 Fig. 1 in page 4 and fig. 2 in page 5
297
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