Journal of South American Earth Sciences 121 (2023) 104165
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Journal of South American Earth Sciences
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Ceratozetidae (Acari: Oribatida) from lower Miocene mexican amber,
including a new species of Trichoribates Berlese, 1910
Margarita Ojeda a, Francisco J. Vega b, Gerardo Rivas a, *
a
b
Facultad de Ciencias, Universidad Nacional Autónoma de México, Coyoacán, Mexico City, 04510, Mexico
Instituto de Geología, Universidad Nacional Autónoma de México, Coyoacán, Mexico City, 04510, Mexico
A R T I C L E I N F O
A B S T R A C T
Keywords:
Acari
Oribatids
Early miocene
Chiapas amber
Mexico
Trichoribates roynortoni Ojeda and Rivas sp. n. Is described based on one adult specimen preserved in estuarine
stratified lower Miocene amber from the Campo La Granja mines, La Quinta Formation, Chiapas Mexico, which
includes mostly estuarine aquatic and semiaquatic animals, but also an important amount of soil arthropods. This
taxon is the first record for the Ceratozetidae family (Acari: Sarcoptiformes: Oribatida) and represents an
addition to the few Acari reported for the Chiapas amber.
1. Introduction
Oribatid mites are an abundant and diverse group in soil environ­
ments around the world. They have an important role in the decaying of
organic matter and nutrient cycling, represented worldwide by 11,435
species (Subías, 2004 updated online 2022).
The fossil record of the Oribatida is one of the largest and oldest
within mites with 70 species known from different types of fossiliferous
deposits, such as limestone and amber. The oldest known representants
of this group correspond to Protochthonius gilboa Norton and Poinar,
1993 and Devonacarus sellnicki Norton and Poinar, 1993 both from the
Middle Devonian of Gilboa, New York, in the United States of America.
Other records correspond to the Upper Jurassic, Cretaceous and Ceno­
zoic (Norton and Poinar, 1993).
Recent reports of Palaeozoic oribatids are those of Subías and Arillo
(2002), who described seven new genera and species: Ctenacaronychus
nortoni Subías and Arillo (2002), from the Upper Devonian in the South
Mountains in USA (New York) and six species from the Lower Carbon­
iferous of Northern Ireland. Arillo et al. (2020) described Epieremulus
sidorchukae Arillo et al., 2020 from Cretaceous (Albian-Cenomanian)
amber of Spain.
The family Ceratozetidae currently has 33 genera and 335 species
(Subías, 2004 updated online, 2022), among them the genus Trichor­
ibates is one of the most diverse, with 64 species recognized. Trichoribates
was established by Berlese (1910) with Murcia trimaculata Koch and
Berendet, 1854 as type species. Weigmann and Norton (2009) discussed
the validity of this species and interpreted the type of Trichoribates, being
one of the most common and taxonomically diverse of oribatid mites.
According to Subías (2004 updated 2022) Trichoribates is represented by
five subgenera: Trichoribates (Trichoribates), T. (Laminizetes), T. (Lat­
ilamellobates), T. (Sacculoribates) and T. (Viracochiella). However, taxo­
nomic placement of some of these taxa have been questioned by
Bayartogtokh and Schatz (2008) and by Behan-Pelletier and Ermilov
(2019). Extant species of Trichoribates are distributed mostly in the
Northern hemisphere, 35 species from the Palaearctic and 17 from the
Nearctic region. Only three species are found in the Neotropics
(T. hammerae Subías, 2004 from Peru, T. serratus Pérez-Ínigo and
Pérez-Ín;igo Jr., 1993 from Brazil, and T. sidorchukae Behan-Pelletier
and Ermilov, 2019 from Ecuador). Behan-Pelletier and Schatz (2010)
mentioned that Ceratozetoidea are found in forest, grassland, tundra and
semi-aquatic habitats. They are found in all biogeographic regions, but
are most diverse at mid to high latitudes in the Nearctic, Palearctic, and
Neotropics. Several Trichoribates species seem to be restricted to harsh
environments with short vegetation periods and show adaptations such
as cold hardiness and prolongation of life cycle (Schatz, 2020).
Fossil records of the family include 20 species in 8 genera from
Paleocene and Quaternary from diverse localities (Dunlop et al., 2020).
There are reports from the Paleogene Baltic amber, with genera such as
Melanozetes and Sphaerozetes: M. foderatus Sellnick (1931);
M. mollicommus fossilis Sellnick (1931); Sphaerozetes convexulus (Koch
and Berendet, 1854) and S. primus Sellnick (1931), being the oldest
species for the group (Koch and Berendet, 1854; Sellnick, 1931).
* Corresponding author.
E-mail address: gerardorivas@ciencias.unam.mx (G. Rivas).
https://doi.org/10.1016/j.jsames.2022.104165
Received 28 September 2022; Received in revised form 16 December 2022; Accepted 16 December 2022
Available online 17 December 2022
0895-9811/© 2022 Elsevier Ltd. All rights reserved.
M. Ojeda et al.
Journal of South American Earth Sciences 121 (2023) 104165
Particularly, subfossils (remains of once living organism with an
incomplete fossilization process) of Trichoribates include five species
from the Quaternary of the Palearctic region: T. biarea Gjelstrup and
Solhøy, 1994; T. incisellus (Koch and Berendet, 1854); T. monticola
(Travé and Vachon, 1975); T. setiger (Trägardh, 1910) and
T. trimaculatus ( Koch and Berendet, 1854) (Dunlop et al., 2020).
Trichoribates is represented by two living species in Mexico: Tri­
choribates (Trichoribates) oztotlicus Palacios-Vargas and Norton (1984)
and T. (Latilamellobates) tepetlensis Palacios-Vargas and Norton (1984),
both described from the Popocatépetl Volcano, whose distribution
shows the Nearctic pattern associated with the largest number of known
species in the northern hemisphere. In lower Miocene (23 Ma) Chiapas
amber, 13 species of mites have been recorded to date (Woolley, 1971;
Norton and Poinar, 1993; Rivas et al., 2016; Rivas and Vega, 2021), all
representatives of the Prostigmata and Oribatida clades. Seven of these
species belong to Oribatida families: Oppiidae, Neoliodidae, Scuto­
verticidae, Mochlozetidae and Oripodidae (Norton and Poinar, 1993)
(Table 1).
As mentioned before, controversy on the recognized generic name,
and the type species, Sphaerozetes (Trichoribates) berleseiJacot, 1929 has
been under discussion, as well as some placement of other genera.
Herein we follow the diagnostic features and classification stated by
Bayartogtokh and Schatz (2008) and Behan-Pelletier and Ermilov
(2019).
The first record of the family Ceratozetidae, subfamily Trichor­
ibatinae for the lower Miocene amber from Mexico is presented in this
paper, with the description of a new species.
Fig. 1. Location map of Simojovel and mines of Campo La Granja, where the
amber piece with the Acari here reported was collected. Modified from Serra­
no-Sánchez et al. (2015).
2. Study area, stratigraphy, paleoenvironment and methods
The lower Miocene amber from Chiapas, Mexico is famous world­
wide for its contents of abundant biological inclusions, among which the
most common are terrestrial insects, followed by arachnids, plants and
occasional fungi (Solórzano-Kraemer, 2007). Aquatic organisms have
also been reported, including algae, ferns, crustaceans, and mollusks
(Serrano-Sánchez et al., 2015). Several amber mines are found near the
town of Simojovel, located in the Sierra Madre de Chiapas (Fig. 1). The
amber-bearing stratigraphic sequence includes three lithostratigraphic
units: the upper portion of La Quinta Formation (Oligocene-Miocene),
the Mazantic Shale and Balumtun Formation (lower Miocene) (Fig. 2).
Isotopic and biostratigraphic data suggest that the amber deposit begun
22.8 Ma ago (Vega et al., 2009; Perrilliat et al., 2010; Sol­
órzano-Kraemer, 2010), in an estuarine environment, nearby the ancient
Gulf of Mexico coast (Serrano-Sánchez et al., 2015). Stratified amber
pieces are found embedded into the hard, calcareous sandstones of the
lower Miocene Finca Carmitto Member of the La Quinta Formation,
from where a diverse estuarine microcrustacean assemblage has been
reported (Serrano Sánchez et al., 2015, among others). Thin sand layers
in amber pieces from Campo La Granja mines (Finca Carmitto Member)
separate what seems to be cyclic events of high tides which flooded the
estuarine floor, leaving planktic and benthic microcrustaceans that got
trapped into small ponds (along with organic matter and other conti­
nental arthropods, including mites), which slowly lose the water due to
exposition of tropical conditions, making the originally liquid
Fig. 2. Stratigraphic section of the Simojovel area, with position of amberbearing sandstone strata from the upper portion of La Quinta Formation
(Finca Carmitto Member). Red stars indicate amber occurrences. Modified from
Serrano-Sánchez et al. (2015).
Table 1
List of fossil Oribatida from Mexican amber, compiled from Woolley (1971);
Norton and Poinar (1993).
Family Neoliodidae
Family Mochlozetidae
Family Oppiidae
Family Oripodidae
Family Scheloribatidae
Family Scutoverticidae
Liodes brevitarsus (Woolley, 1971)
Mochloribatula smithi (Woolley, 1971)
Oppia mexicanus (Woolley, 1971)
Oppia setiger (Woolley, 1971)
Benoibates chiapanensis (Woolley, 1971)
Parapirnodus denaius (Woolley, 1971)
Scheloribates durhami (Woolley, 1971)
Arthrovertex hurdi (Woolley, 1971)
resin/water environment into a viscous and eventually solid resin blocks
that later were buried and became part of the sandstone sequence of the
Finca Carmitto Member. Compared to the relatively soft Mazantic Shale
amber, the Campo La Granja amber blocks are very hard to cut and
polish due to the abundant contents of sandstone.
The specimens here reported were originally observed under an
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Journal of South American Earth Sciences 121 (2023) 104165
American Optical stereo microscope. The amber piece was sectioned
with a small diamond saw to get tiny pieces which were polished with
sandpaper to reach an approximate thickness of about 3 mm. The pieces
were polished with car polish and final details were obtained with
Brasso®. Once the small pieces were completely translucid, specimens
were observed and photographed with a stereo microscope Zeiss Axio
Zoom v16, to obtain several images that were processed with Helicon
Focus to obtain high resolution images. Subsequently, the tiny pieces
were reduced to slim and circular plates of 1.5 cm in diameter, according
to the method of Sidorchuk and Vorontsov (2018) for a more detailed
observation in a compound microscope. Images were obtained using an
AxioCam MRC5 camera using a Carl Zeiss Axio Zoom microscope. From
these and with the help of an optical microscope, the details of the
morphology were compared, and later with the CorelDraw Graphic Suite
program (version 2020) the corresponding drawings were made. The
identification was carried out using Balogh (1972), Balogh and Balogh
(1992), Bayartogtokh and Schatz (2008), and Behan-Pelletier and
Ermilov (2019).
The terminology used follows Grandjean (for references see Travé
and Vachon, 1975), and Norton and Behan-Pelletier (2009). The
following measurement and notational conventions are used:
lam—lamela; tlam—translamela; lpr –lateral prodorsal ridge, tu—tuto­
rium; gt—genal tooth; ro, le, in, Ss, ex—rostral, lamelar, interlamelar,
bothridial and exobothridial setae, respectively; bo—botridio;
len—lenticulus; Ad—dorsosejugal porose area; A3— notogastral porose
area; Pd I, Pd II—pedotectum I and II, respectively; cus—custodium;
dis—discidium; cp—circumpedal carina; pg—genital shield; Tr, Fe, Ge,
Ti, Ta—trochanter, femur, genu, tibia and tarsus of legs, respectively; σ,
φ—solenidia of legs; setae of legs—v, ev, bv, l, d, ft, tc, it, p, u, a, s, pv.
3. Results
3.1. Systematic Palaeontology
Fig. 3. Trichoribates roynortoni Ojeda and Rivas sp. n. A. and B. dorsal view of
body. C. Prodorsum showing rostrum, tutoria and lamellar setae. D. Lamellar
cusps. E. Shape of prodorsal setae: le (lamellar), ro (rostral), bs (bothridial or
Sensilli). F. Solenidia of genua, tibia and tarsus of leg I. Holotype IGM-13079.
Subclass Acari Leach, 1817.
Superorder Acariformes Zachvatkin, 1952.
Order Sarcoptiformes Reuter, 1909.
Suborder Oribatida Dugès, 1834.
Family Ceratozetidae Jacot, 1925.
Genus TrichoribatesBerlese, 1910 .
Trichoribates roynortoni sp. n. Ojeda and Rivas, 2022.
Type specimen
Holotype IGM-13079, deposited at the Colección Nacional de Pale­
ontología, Instituto de Geología, Universidad Nacional Autónoma de
México, Mexico City 04510, Mexico.
Etymology
The name of the species is dedicated to Dr. Roy A. Norton, one of the
most outstanding acarologists, teacher, mentor and colleague to many.
This species is assigned in homage to his brilliant career, which includes
multiple works to discover oribatid fossils.
3.2. Description
3.2.1. Size - dimensions
Total length (from the end of the rostrum to the edge of the
notogaster) 907 μm. Prodorsum 148 μm long and 256 μm wide (at the
level of the sejugal suture). Notogaster: length 769 μm, and 680 μm
width (at the level of coxa IV).
Fig. 4. Trichoribates roynortoni Ojeda and Rivas sp. n. A. habitus ventral, B.
genital region. C. prodorsal area D. Lamella. Holotype IGM-13079.
3.2.2. Integument
Deep dark brown, smooth and with shiny appearance (Figs. 3A and
4). In part of the edge of the prodorsum and the region of the pter­
omorphs, a slight roughness is appreciated (Fig. 3B).
rounded, ending in a small bulge in the middle, without incisions, teeth
or grooves (Figs. 3C and 4C, D). Tutoria present, slightly wide lamelli­
form parallel to the dorsal edge of prodorsum and lamellae, extending
anteriorly to the insertion of the ro setae, triangular in its distal portion
(Fig. 3D). Lamellae wide and long, in the distal part the well-developed
cusps are observed from which the lamellar setae emerges (le, 109 μm),
3.2.3. Prodorsum
Triangular shape (almost of an equilateral triangle). Rostrum
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Journal of South American Earth Sciences 121 (2023) 104165
cusps (length similar to the 1:1 lamellae) with a pair of “teeth” (dens),
the external one longer than the internal. Lamellae tapered distally and
slightly converge (Fig. 3D). Translamella narrow and long (approxi­
mately 100 μm between lamellae), straight (the distance between the
lamellae corresponds to the length of the cusps) and forms a trapezoidal
area between the lamellae (Fig. 3B). Prodorsal setae barbed in their
distal half. Rostral setae (ro, 98 μm) emerge at level of the lamellae from
the edge of the prodorsum. Interlamellar setae (in, 197 μm), almost
twice as long as the lamellar setae (le), they are the longest in the pro­
dorsum and are inserted almost on the edge of it, close to the sejugal
suture. Small bothridial setae (Sensilli) (69 μm) with a clavate head and
a long pedicel, apparently covered by the anterior edge of notogaster
(Fig. 3E).
second specimen, some details of their morphology are also observed
that allow it to be assigned to the Ceratozetidae family; however, several
of the characters necessary for the generic and specific determination
cannot be analyzed.
4. Remarks
We observe some of the morphological features stated in the revised
diagnosis (Bayartogtokh and Schatz, 2008; Behan-Pelletier and Ermilov,
2019) for the genus Trichoribates sensu stricto. This included the shape of
rostrum rounded; lamella wide, with cusp and translamella; lamellar
cusp with lateral and median dens; bothridium cup-shaped; sensillus
with an oval head (these two last characteristics observed under mi­
croscope). Tutorium broad, with cusp pointed distally. Notogaster with
large pteromorph curved ventrally; lenticulus present as a small round
area. Notogastral setae not visible; of the four pairs of notogastral porose
areas, A2 and A3 seem to be present. Legs heterotridactylous; tibia I with
dorsal distal apophysis bearing solenidion φ2 and seta l” of tibiae and
genua I, II and sometimes that of tibiae and genua III, IV thick, heavily
barbed.
The new species can be separated from the three extant, described for
the Neotropical Region (and the two described for Mexico) by the
following characteristics: Trichoribates roynortoni sp. n. is the largest
species with 900 μm, the closest species in terms of size are those
described for the Popocatépetl Volcano, Trichoribates (Latilamellobates)
tepetlensis Palacios-Vargas and Norton, 1984 (739–854 μm) and
T. (Trichoribates) oztotlicus Palacios-Vargas and Norton, 1984 (624–768
μm), so we can point out that the Mexican species are among the largest
in terms of size.
The rostrum is smooth and ends in a rounded vertex, as seen in
T. hammerae Subías, 2004 and T. sidorchukae Behan-Pelletier and
Ermilov (2019). The shape of the lamellae and translamella is very
similar to that described for T. serratus Pérez-Iñigo and Pérez-Iñigo Jr.
(1993) but differs from it in the shape and size of the pteromorphs, since
in T. roynortoni sp. n. they are more rounded in their distal part.
T. serratus is a small species compared to T. roynortoni sp. n. (it only
measures about half: 473 vs 907 μm). Additionally, both lateral and
medial teeth of cusp are present as in T. serratus and T. hammerae Subías,
2004. The size of the cusps with the external (lateral) and internal
(medial) teeth are approximately the same length (1:1.1 to 1.2) as the
lamellae; in T. serratus Pérez-Iñigo and Pérez-Iñigo Jr. (1993) the
lamellae are longer (1:1.8), as well as in T. sidorchukae Behan-Pelletier
and Ermilov (2019), where the lamellar cusps are longer than the length
of lamella. The translamella is straight, the distance between the
lamellae corresponds to the length of the cusps, unlike the species
recorded in the Neotropics, not in a “V" or “U" shape but rather a trap­
ezoidal area between the lamellae (Fig. 3A and B). The body is pear
shaped, slightly narrower in the anterior region.
Comparison of T. roynortoni sp. n. with some of the descriptions of
the subfossil species known to date, corroborates its inclusion within the
genus. As it happens with the extant species with distribution in the
neotropics, regarding the size the new species is the largest among
T. biarea Gjelstrup and Solhøy, 1994 (550–600 μm); T. incisellus (Koch
and Berendet, 1854) (550 μm), T. monticola (Travé and Vachon, 1975)
(510 μm), T. setiger (Trägardh, 1910) (495 μm). All the aforementioned
species have a Palearctic distribution.
Regarding the shape of lamellae and translamellae, in T. biarea,
T. incisellus and T. roynortoni sp. n., lamellae have cusp with outer dens
larger than the inner one; as for the translamellae both T. biarea and
T. roynortoni sp. n. Are similar, narrow, as wide as long, with a U shape.
With T. monticola, the new species shares the shape of the translamellae
(a wider U shape), but differs from this because lamellae do not have
cusp and are wider at their distal portion.
3.2.4. Notogaster
Rounded, pear-shaped, longer than wide (769 × 680 μm). Smooth
cuticle, some areas with slight striae (Fig. 3A). Sejugal suture complete
and rounded, in shape of an inverted and wide “U". Notogastral setae not
evident. Presence of porous areas only observed on the posterior border
— A3 as a lighter area on the posterior border of the notogaster (and not
clearly, Fig. 3A). Humeral extension present. Lenticulus poorly devel­
oped (Figs.. 3A, 4C-D), shown as a less pigmented area. Pteromorphs
well developed, small immobile-type slightly curved ventrally, without
an obvious insertion area between them and notogaster.
3.2.5. Epimeral region
Typical of oribatid poronotic. Genital plate round, genital setae not
evident (Fig. 4A–B). Apodemes well developed. Epimeral setae not
distinguishable.
3.2.6. Legs
Median claws thicker than the lateral ones. Legs I-IV subequal in
length, elongated, cylindrical. Solenidium ⱳ1 on tarsi I slightly longer
than ⱳ2. Solenidium φ1 arises from a well-developed tibial process
(Fig. 3F). Seta l′′ of genua I and II setiform (Fig. 3F). Tibia I with a
relatively small apophysis from which the solenidium φ2 emerges. Sol­
enidium φ1 of tibia I twice the length of φ2. Genua I with ventro-distal
projection. Most setae barbed; some smooth, such as the setae (p) and (u)
of the tarsus I–IV.
An additional oribatid mite IGM-13080 (Fig. 5), located close to the
previous one, was analyzed in the same amber piece (<5 mm). For this
Fig. 5. Specimen of Ceratozetidae. A and B dorsal aspects of the body. C.
Rostrum. D. Leg III. Specimen IGM-13080.
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Journal of South American Earth Sciences 121 (2023) 104165
5. Conclusions
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Oribatid mites are one of the groups with a large and ancient fossil
record with over 70 species. Amber deposits in the state of Chiapas have
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1993). Several species of the family Ceratozetidae have been recorded as
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CRediT authorship contribution statement
Margarita Ojeda: Writing – review & editing, Writing – original
draft, Visualization, Validation, Supervision, Methodology, Investiga­
tion, Formal analysis, Data curation. Francisco J. Vega: Writing – re­
view & editing, Writing – original draft, Validation, Supervision, Project
administration, Methodology, Investigation, Formal analysis, Data
curation, Conceptualization. Gerardo Rivas: Writing – review & edit­
ing, Writing – original draft, Validation, Methodology, Investigation,
Formal analysis, Data curation, Conceptualization.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
No data was used for the research described in the article.
Acknowledgments
We thank Biol. Susana Guzmán, Laboratorio de Microscopía y
Fotografía 2-LaNaBio, Instituto de Biología, UNAM for her help with
some of the microphotographs, to Violeta Romero Mayén (Colección
Nacional de Paleontología, Instituto de Geología, UNAM), for her help in
assigning collection numbers, and to Aurora Vassallo for her help with
the figure plates. Special mention to Dr. Roy A. Norton and Dr. Valerie
Behan-Pelletier for their invaluable help by providing us with some of
the literature of Oribatida and the comments on the taxonomic place­
ment of the subfossil specimens. The authors appreciate the constructive
comments on this paper given by Dr. Antonio Arillo, and an anonymous
reviewer who help to improve the original manuscript.
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Journal of South American Earth Sciences 121 (2023) 104165
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