ARTHROPODS OF LA SELVA (PROJECT ALAS): ORIBATIDA

ARTHROPODS OF LA SELVA (PROJECT ALAS): ORIBATIDA
Valerie Behan-Pelletier

Biodiversity Program, ECORC, Research Branch, Agriculture and Agri-Food Canada, K. W. Neatby Bldg., Ottawa, Ontario, Canada K1A 0C6.

Email: behanpv@em.agr.ca

http://res.agr.ca/ecorc/staff/beha-v.htm

Objectives
Personnel and infrastructure
Approach
Results
Taxonomic list of Oribatida of La Selva
Collaborators
Discussion
References
Comments on some oribatid species from La Selva

OBJECTIVES

The systematics and ecology of the brachypyline oribatid mite fauna of primary and secondary lowland tropical rainforest are the subject of ongoing research as part of the Arthropods of La Selva (ALAS) Project (Colwell 1996). Estación Biologica La Selva at 10°26’N 84°1’W and 50-150m elevation, is the biotically rich field station of the Organization for Tropical Studies in the Atlantic lowland, evergreen tropical rainforest of Costa Rica (McDade et al. 1993). Prior to the ALAS Project little was known of the oribatid fauna of Costa Rica (Balogh and Balogh 1988, 1990; Balogh 1994).

Project ALAS addresses a fundamental question: How many arthropod species are there in a tropical rainforest? To answer this question systematists generally use a "find them all" approach, using specialized knowledge of the ecology and behavior of particular taxonomic groups to discover many individuals of those groups efficiently. In contrast, ecologists use a "sample and estimate" approach. By taking quantitative samples and making assumptions about community structure, they can make statistical estimates of species richness. Project ALAS combines both approaches in an assessment of arthropod diversity. "Sample and estimate" methods are used to sample a set of broad "survey taxa," while "find them all" methods are used to sample much more thoroughly a set of smaller "focal taxa." The known focal taxa are then used to calibrate, and evaluate and compare the sampling and estimation methods.

PERSONNEL AND INFRASTRUCTURE

The core of Project ALAS is the four parataxonomists (pictured above) who are resident near La Selva and an on-site air-conditioned laboratory with collection facilities (Figure), such as Berlese extractors, microscopes, specimen bar-coding technology, sophisticated data capture and database management using BIOTA (Colwell 1997), and reliable email and internet connectivity.

The parataxonomists have been trained in quantitative and qualitative collecting techniques for oribatid mites including soil, litter and fungal sampling and extraction using Berlese funnels, twig and leaf washing techniques and heptane flotation, and canopy fumigation. They are trained at recognizing and sorting the major oribatid cohorts and in recognizing members of the brachypyline Oribatida to morphogenus and morphospecies. They are also trained in the preparation of slide mounted specimens, in preparation and maintenance of an alcohol collection of Oribatida, and in use of the biodiversity data management program BIOTA.

APPROACH

Quantitative samples from litter and surface soil in primary and secondary rainforest were collected monthly for 12 months during 1993. Mites were extracted using Berlese funnels based on the design of Norton and Kethley (1988) (Figure) and brachypyline oribatid mites sorted to morphogenus and morphospecies. Quantitative canopy fumigation was carried out during the same period and brachypyline Oribatida were also sorted from these samples.

During 1994-1996 additional Berlese samples were collected at Estación Biologica La Selva from a variety of habitats in addition to litter and surface soil, including decaying wood, fungi, leaves, flowers and semiaquatic habitats. Twig washing with subsequent heptane flotation was carried out on samples taken from the canopy of a number of tree species. The detailed habitat data associated with these collections is recorded in BIOTA and is easily accessible.

The ALAS staff selected specimens and sorted from these sample units. They did not sort every oribatid mite from a sample, rather they selected up to 5 specimens of everything that looked like a distinct brachypyline species. Undoubtedly there are splitting errors and lumping errors, but clearly this was a more efficient and taxonomically valuable protocol than either slide mounting every oribatid in a sample, or expecting all taxonomists to sort through material for the taxa of their interest.

Ceratozetoidea were one of the focal groups of Oribatida in Project ALAS, with brachypyline Oribatida being the survey taxon. Brachypylina were chosen as the survey taxon rather than all Oribatida, because the former is monophyletic, and because oribatid numbers and diversity in any soil and litter sample at La Selva can be overwhelming. However, because of their ease of recognition the families Lohmanniidae, Crotoniidae and Phthiracaridae were also sorted to morphospecies. Among the Brachypylina the most diverse families at Estación Biologica La Selva are the Oppiidae, Microzetidae, Scheloribatidae, Haplozetidae, and Galumnidae

RESULTS

As a result of this collecting and sorting effort the oribatid collections associated with Project ALAS are extensive, and include the following:

approximately 600 vials containing mites in ETOH sorted to named species, or sorted and identified to genus (Table 1);
approximately 1600 vials containing an average of 5 mites/vial of Oppioidea in ETOH, sorted to morphogenus, but not identified;
approximately 200 vials containing Oripodoidea, other than Mochlozetidae, in ETOH, sorted to morphospecies, but not indentified;
approximately 700 slides primarily of brachypyline Oribatida, identified as far as possible (Table 1);
residues for all collections which can be accessed for further specimens of specific taxa, or immatures, or specimens of Macropylina.

Table 1. Oribatida of Estación Biologica La Selva

Superfamily	Family	Genus	Species	Author
Brachychthonioidea	Brachychthoniidae	Brachychthonius	at least 2 spp.
Hypochthonioidea	Hypochthoniidae		sp.
Hypochthonioidea	Hypochthoniidae	Eohypochthonius	at least 2 spp.
Hypochthonioidea	Mesoplophoridae	Mesoplophora	nr. longiseta	Mahunka 1982
Hypochthonioidea	Mesoplophoridae		at least 2 spp.
Epilohmannioidea	Epilohmanniidae	Epilohmannia	sp.
Lohmannioidea	Lohmanniidae		at least 3 spp.
Lohmannioidea	Lohmanniidae	Javacarus	porosus	Hammer 1980
Phthiracaroidea	Phthiracaridae	Hoplophorella	at least 3 spp.
Phthiracaroidea	Phthiracaridae	Hoplophthiracarus	nr. latebrosus	Niedbala 1982
Phthiracaroidea	Phthiracaridae	Hoplophthiracarus	at least 2 other spp.
Phthiracaroidea	Phthiracaridae	Austrophthiracarus	nr. dubius	Niedbala 1982
Phthiracaroidea	Phthiracaridae	Archiphthiracarus	sp.
Phthiracaroidea	Phthiracaridae		at least 3 other spp.
Euphthiracaroidea	Euphthiracaridae	Brasiliotritia	brasiliensis	Maerkel 1964
Euphthiracaroidea	Euphthiracaridae	Rhysotritia	comteae	Mahunka 1983
Crotonioidea	Crotoniidae	Crotonia	sp.
Malaconothroidea	Trhypochthoniidae		sp.
Malaconothroidea	Trhypochthoniidae	Afronothrus	sp.
Malaconothroidea	Trhypochthoniidae	Allonothrus	neotropicus	Balogh & Mahunka 1969
Malaconothroidea	Trhypochthoniidae	Archegozetes	sp.
Malaconothroidea	Malaconothridae		sp.
Malaconothroidea	Malaconothridae	Malaconothrus	at least 2 spp.
Nanhermannioidea	Nanhermanniidae		sp.
Nanhermannioidea	Nanhermanniidae	Cyrthermannia	nr. guadeloupensis	Mahunka 1985
Hermanniellioidea	Hermanniellidae	Ampullobates	sp.
Hermanniellioidea	Hermanniellidae		at least 3 other spp.
Hermanniellioidea	Plasmobatidae		sp.
Hermanniellioidea	Plasmobatidae	Orbiculobates	sp.
Hermanniellioidea	Plasmobatidae	Plasmobates	sp.
Hermanniellioidea	Plasmobatidae	Solenozetes	sp.
Liodoidea	Liodidae	Teleioliodes	sp.
Liodoidea	Liodidae	Teleioliodes	madininensis	Grandjean 1934
Liodoidea	Liodidae	Liodes	nr. zikani	(Sellnick 1930)
Plateremaeoidea	Pedrocortesellidae	Pedrocortesella	sp.
Plateremaeoidea	Pheroliodidae	Pheroliodes	sp.
Cepheoidea	Microtegeidae	Microtegeus	at least 2 spp.
Cepheoidea	Microtegeidae	Microtegeus	borhidii	Balogh & Mahunka 1974
Cepheoidea	Microtegeidae	Microtegeus	nr. similis	Balogh & Mahunka 1980
Cepheoidea	Eremaeozetidae	Eremaeozetes	nr. ursulae	Mahunka 1985
Cepheoidea	Eremaeozetidae	Eremaeozetes	at least 5 other species
Microzetoidea	Microzetidae	Berlesezetes	sp.
Microzetoidea	Microzetidae	Berlesezetes	auxiliaris	(Grandjean 1936)
Microzetoidea	Microzetidae	Licnozetes	flabellatus	Balogh & Mahunka 1969
Microzetoidea	Microzetidae	Rugozetes	gladiator	Balogh 1962
Microzetoidea	Microzetidae	Schalleria	sp.
Microzetoidea	Microzetidae	Stylozetes	sp.
Microzetoidea	Microzetidae	Undolozetes	sp.
Microzetoidea	Microzetidae		at least 4 other spp.
Amerobelboidea	Eremulidae	? genus.	sp.
Amerobelboidea	Eremulidae	? Eremulus	sp.
Amerobelboidea	Eremulidae	Eremulus	at least 2 spp.
Amerobelboidea	Eremulidae	Eremulus	rigidisetosus	Balogh & Mahunka
Amerobelboidea	Eremulidae	Eremulus	cf. translamellatus	Balogh & Mahunka
Amerobelboidea	Damaeolidae	Caudamaeolus	sp
Amerobelboidea	Heterobelbidae	Heterobelba	cf. crassisetosa	Beck 1962
Gustavioidea	Astegistidae	Cultroribula	at least 2 spp.
Gustavioidea	Xenillidae	Xenillus	nr. venezuelanus	Balogh & Balogh 1984
Gustavioidea	Xenillidae	Xenillus	at least 2 other spp.
Gustavioidea	Peloppiidae	Ceratorchestes	cornutus	Mahunka 1982
Gustavioidea	Peloppiidae	Ceratorchestes	nr. baloghi	Mahunka 1982
Gustavioidea	Peloppiidae	Ceratorchestes	at least 2 other spp.
Tectocepheoidea	Tectocepheidae	Tectocepheus	sp.
Tectocepheoidea	Tectocepheidae	Tegeozetes	sp.
Tectocepheoidea	Tectocepheidae	Tegeozetes	tunicatus	Berlese 1913
Carabodoidea	Carabodidae	Carabodes	nr. octogonalis	Balogh & Mahunka 1969
Carabodoidea	Carabodidae	Neocarabodes	sexpilosus	Balogh & Mahunka 1969
Carabodoidea	Carabodidae	Spathulocepheus	sp.
Carabodoidea	Carabodidae	Yoshiobodes	sp.
Carabodoidea	Otocepheidae	Dolicheremaeus	bolivianus	Balogh & Mahunka 1969
Carabodoidea	Otocepheidae	Dolicheremaeus	at least 4 other species
Oppioidea	Dampfiellidae	Beckiella	sp.
Oppioidea	Oppiidae	? Genus	sp.
Oppioidea	Oppiidae	? Genus	sp.
Oppioidea	Oppiidae	Amerioppia	sturmi	Balogh 1984
Oppioidea	Oppiidae	Amerioppia	sp. 2
Oppioidea	Oppiidae	Brachyoppiella	sp.
Oppioidea	Oppiidae	Pulchroppiella	flagellata	Mahunka 1983
Oppioidea	Oppiidae	Pulchroppiella	sp.
Oppioidea	Oppiidae	Striatoppia	sp
Oppioidea	Oppiidae	Globoppia	trichotos	Balogh & Mahunka 1977
Oppioidea	Oppiidae	Karenella?	at least 2 spp.
Oppioidea	Oppiidae	Lanceoppia	at least 2 spp.
Oppioidea	Oppiidae	Teratoppia	nr. centroamericana	Woas 1986
Oppioidea	Oppiidae	Varioppia	sp.
Oppioidea	Oppiidae		at least 15 other spp.
Oppioidea	Granuloppiidae	Bornemiszaella	at least 3 spp.
Oppioidea	Sternoppiidae	Sternoppia	sp.
Oppioidea	Suctobelbidae		at least 3 spp.
Oppioidea	Rhynchoribatidae	Rhynchoribates	at least 2 spp.
Ameronothroidea	Tegeocranellidae	Tegeocranellus	alas	Behan-Pelletier 1997
Cymbaeremaeoidea	Cymbaeremaeidae	Scapheremaeus	at least 4 species
Licneremaeoidea	Licneremaeidae	Licneremaeus	sp.
Licneremaeoidea	Scutoverticidae	Arthrovertex	at least 2 spp.
Oripodoidea	Mochlozetidae	? Genus	sp.
Oripodoidea	Mochlozetidae	Mochloribatula	sp.
Oripodoidea	Mochlozetidae	Mochlozetes	sp.
Oripodoidea	Mochlozetidae	Uracrobates	sp.
Oripodoidea	Mochlozetidae		at least 4 other spp.
Oripodoidea	Xylobatidae	Brasilobates	at least 2 spp.
Oripodoidea	Xylobatidae	Xylobates	at least 5 spp.
Oripodoidea	Protoribatidae	Liebstadia	at least 2 spp.
Oripodoidea	Protoribatidae	Maculobates	sp.
Oripodoidea	Oribatulidae	? genus	sp.
Oripodoidea	Oribatulidae	? Genus	sp.
Oripodoidea	Oribatulidae	Cordiozetes	sp.
Oripodoidea	Oribatulidae	Phauloppia	at least 2 spp.
Oripodoidea	Oribatulidae	Phylloribatula	sp.
Oripodoidea	Neotrichozetidae	Neotrichozetes	sp.
Oripodoidea	Haplozetidae	Haplozetes	nr. minimicoma	Beck 1964
Oripodoidea	Haplozetidae	Haplozetes	triangulatus	Beck 1964
Oripodoidea	Haplozetidae	Haplozetes	sp. 2
Oripodoidea	Haplozetidae	Peloribates	at least 2 spp.
Oripodoidea	Haplozetidae	Rostrozetes	ovulum	(Berlese 1908)
Oripodoidea	Haplozetidae	Rostrozetes	cf. geminisetosus	Balogh & Mahunka 1978
Oripodoidea	Haplozetidae	Rostrozetes	glaber	Beck 1965
Oripodoidea	Haplozetidae	Rostrozetes	dimorphochaites	Higgins 1966
Oripodoidea	Haplozetidae	Rostrozetes	cf. carinatus	Beck 1965
Oripodoidea	Haplozetidae	Rostrozetes	cf. ovulum
Oripodoidea	Haplozetidae	Rostrozetes	at least 5 other species
Oripodoidea	Nasobatidae	Nasobates	sp.
Oripodoidea	Scheloribatidae	? genus	sp.
Oripodoidea	Scheloribatidae	Cryptozetes	at least 2 spp.
Oripodoidea	Scheloribatidae	Grandjeanobates	sp.
Oripodoidea	Scheloribatidae	Ischeloribates	at least 3 spp.
Oripodoidea	Scheloribatidae	Oxyscheloribates	sp.
Oripodoidea	Scheloribatidae	Scheloribates	cf. bidactylus	Hammer 1961
Oripodoidea	Scheloribatidae	Scheloribates	nr. praeincisus rotundiclava	Perez-Inigo & Baggio 1986
Oripodoidea	Scheloribatidae	Scheloribates	at least 13 other spp.
Oripodoidea	Scheloribatidae	Stelechobates ?	sp.
Oripodoidea	Oripodidae	? genus	sp.
Oripodoidea	Oripodidae	? genus	sp.
Oripodoidea	Oripodidae	Benoibates	at least 2 spp.
Oripodoidea	Oripodidae	Cordiozetes	sp.
Oripodoidea	Oripodidae	Exoribatula	sp.
Oripodoidea	Oripodidae	Oripoda	nr. lobata	Mahunka 1985
Oripodoidea	Oripodidae	Oripoda	at least 2 other spp.
Oripodoidea	Oripodidae	Truncopes	sp.
Oripodoidea	Parakalummidae	Neoribates	nr. erectus	(Balogh & Mahunka 1969)
Oripodoidea	Parakalummidae	Neoribates	at least 2 other spp.
Oripodoidea	Parakalummidae	Parakalumma	foveolata	Balogh & Mahunka 1969
Ceratozetoidea	Ceratozetidae	Selvazetes	sylvanus	Behan-Pelletier 1998
Ceratozetoidea	Ceratozetidae	Ceratozetes	ambiguus	Behan-Pelletier 1998
Oripodoidea	Ceratozetidae	Ceratozetes	nr. platyrhinus	Hammer
Ceratozetoidea	Mycobatidae	Allozetes	lacandonicus	Mahunka & Palacios-Vargas 1996
Ceratozetoidea	Mycobatidae	Allozetes	alas	Behan-Pelletier 1998
Ceratozetoidea	Mycobatidae	Lamellobates	intermedius	Nübel-Reidelbach & Woas 1992
Ceratozetoidea	Mycobatidae	Lamellobates	reticulatus	Behan-Pelletier 1998
Ceratozetoidea	Mycobatidae	Paralamellobates	striatus	Behan-Pelletier 1998
Ceratozetoidea	Mycobatidae	Punctoribates	sp.
Ceratozetoidea	Zetomimidae	Heterozetes	heleios	Behan-Pelletier 1998
Ceratozetoidea	Zetomimidae	Zetomimus	naias	Behan-Pelletier 1998
Oribatelloidea	Oribatellidae	Oribatella	at least 3 spp.
Galumnoidea	Epactozetidae	Epactozetes	sp.
Galumnoidea	Epactozetidae	Truncozetes	sp.
Galumnoidea	Ceratokalummidae	Achipterina	sp.
Galumnoidea	Ceratokalummidae	Arcozetes	sp.
Galumnoidea	Galumnellidae	Galumnopsis	sp.
Galumnoidea	Galumnidae	Galumna	at least 3 spp.
Galumnoidea	Galumnidae	Allogalumna	at least 3 spp.
Galumnoidea	Galumnidae	Pergalumna	at least 8 spp.
Galumnoidea	Galumnidae		at least 4 other species

Total	56	118	264

The information in Table 1 is clearly incomplete but does allow coarse comparison with oribatid species lists from other well-studied sites, such as temperate deciduous forest at the Coweeta LTER (http://sparc.ecology.uga.edu/webdocs/species/.

COLLABORATORS

As indicated in Table 1 most oribatid species at La Selva are undescribed. One of the goals of Project ALAS is to encourage taxonomic specialists to take advantage of the wealth of material that has been collected and sorted, with the associated detailed collection locality data. To date Project ALAS — Oribatida has the following taxonomic collaborators:

TAXONOMIC GROUP	SPECIALIST	ADDRESS
Phthiracaroidea	Wotjek Niedbala	Department of Animal Taxonomy and Ecology A. Mickiewicz University Szamarzewskiego 91A PL 60-569 Poznan Poland niedbala@main.amu.edu.pl
Lohmanniidae	Roy Norton	State University of New York Syracuse, New York USA 13210 ranorton@mailbox.syr.edu
Crotonioidea	Ziemowit Olszanowski	Department of Animal Taxonomy and Ecology A. Mickiewicz University Szamarzewskiego 91A PL 60-569 Poznan Poland olszanow@amu.edu.pl
Scapheremaeus	Matt Colloff	Department of Entomology, CSIRO Canberra Australia matthew.colloff@ento.csiro.au
Eremaeozetidae, Lohmanniidae	Heinz Schatz	Institute of Zoology University of Innsbruck Technikerstrasse 25 A-6020 Innsbruck Austria heinrich.schatz@uibk.ac.at
Ceratozetoidea	Val Behan-Pelletier	Acarology Unit, ECORC K.W. Neatby Bldg. 960 Carling Ave. Ottawa, Ontario, Canada K1A 0C6 behanpv@em.agr.ca
Galumnidae	Sandor Mahunka	Hungarian Natural History Museum Budapest Baross u.13. H-1088 Hungary mahunka@zoo.zoo.nhmus.hu

and Project ALAS encourages further collaboration, with taxonomists borrowing this well-prepared material and incorporating it into their research on specific taxonomic groups. Specimens can be borrowed on very "long-term" loans from Insitituto Nacional de Biodiversidad, Santo Domingo, Costa Rica (INBio). INBio seeks collaboration with taxonomists who will return data about the Costa Rican fauna, contribute expertise and return identified specimens. INBio also has a generous specimen retention policy for specimens that are not unique.

DISCUSSION

The approach used in Project ALAS, i.e., incorporation of detailed specimen and collection data into BIOTA, allows a range of questions to be addressed that are of value to ecologists, including:

habitat associations for each identified species;
microhabitat and seasonality of the most common species;
microhabitat and seasonality of rare species;
common and rare species for primary vs. secondary rainforest;
common and rare species for soil and litter vs. canopy.

A key question posed by Project ALAS is how effective quantitative collecting techniques are in assessing total species richness in any area. Although there are few representatives of Ceratozetoidea at La Selva, they can be used to address this question. Among ceratozetoid species only Selvazetes sylvanus and Lamellobates intermedius were collected in the quantitative Berlese sampling completed in January 1994, and the latter species was collected on only one occasion (Behan-Pelletier 1998). The nine other species in the superfamily (Table 1) were only found as a result of more focussed collecting from specialized habitats, such as the experimental swamp, riparian habitats, and from grasses or the leaves of plants. Thus, only about one-fifth of the ceratozetoid fauna was assessed using standard quantitative sampling methods. Whether the same ratio is applicable to other oribatid families and superfamilies will be addressed by ongoing and future research as part of the ALAS Project.

REFERENCES

Balogh J. and Balogh P. 1988. Oribatid mites of the Neotropical Region I. Akadémiai Kiadó, Budapest. 335pp.

Balogh J. and Balogh P. 1990. Oribatid mites of the Neotropical Region II. Akademiai Kiadó, Budapest: 1-333.

Balogh P. 1994. New Granuloppiidae (Acari: Oribatei, Oppioidea) from Costa Rica. Acta Zool. Acad. Scient. Hung. 40:15-19.

Behan-Pelletier, VM. 1998. Ceratozetoidea (Acari: Oribatida) of lowland tropical rainforest, La Selva, Costa Rica. Acarologia 39: 349-381.

Colwell R. 1996. ALAS Web site: http://viceroy.eeb.uconn.edu/ALAS/ALAS.html.

Collwell R. 1997. Biota: The Biodiversity Database Manager. Sinauer Associates. http://viceroy.eeb.uconn.edu/biota.

McDade LA, Bawa KS, Hespenheide HA, and Hartshorn GS. (Eds.) 1993. La Selva, ecology and natural history of a neotropical rainforest. University of Chicago Press, Chicago, Illinois.

Norton RA. and Kethley J. 1988. A collapsible full-sized Berlese-funnel system. Ent. News 99:41-47.

COMMENTS ON SOME ORIBATID SPECIES FOUND AT ESTACION BIOLOGICA LA SELVA

Haplozetidae: Rostrozetes ovulum (Berlese)
Mycobatidae: Lamellobates reticulatus Behan-Pelletier
Neotrichozetidae: Neotrichozetes
Mycobatidae: Paralamellobates striatus Behan-Pelletier
Mycobatidae: Selvazetes sylvanus Behan-Pelletier
Tegeocranellidae: Tegeocranellus alas Behan-Pelletier
Zetomimidae: Heterozetes heleios Behan-Pelletier
Zetomimidae: Zetomimus naias Behan-Pelletier

Page author: Valerie Behan-Pelletier, Biodiversity Program, ECORC, Research Branch, Agriculture and Agri-Food Canada, K. W. Neatby Bldg., Ottawa, Ontario, Canada K1A 0C6. Email: behanpv@em.agr.ca

Web page assistance: John T. Longino, The Evergreen State College, Olympia WA 98505 USA.longinoj@evergreen.edu

Date of this version: 9 September 1999

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