# Distributions, range extensions and host list update for four phoretic mite genera (Acari: Klinckowstroemiidae) on Passalidae (Coleoptera)

Jack Schuster, Daniel Gygax, Dafna L Diaz, Francisco Serrano-Peraza

► To cite this version:

Jack Schuster, Daniel Gygax, Dafna L Diaz, Francisco Serrano-Peraza. Distributions, range exten-
sions and host list update for four phoretic mite genera (Acari: Klinckowstroemiidae) on Passalidae
(Coleoptera). Acarologia, 2023, 63 (2), pp.596-604. 10.24349/1zn2-hq18 . hal-04114659

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# Distributions, range extensions and host list update for four phoretic mite genera (Acari: Klinckowstroemiidae) on Passalidae (Coleoptera)

Jack Schuster<sup>a</sup>, Daniel Gygax<sup>a</sup>, Dafna L. Diaz<sup>b</sup>, Francisco Serrano-Peraza<sup>b</sup>

## Original research

### ABSTRACT

Mites of the family Klinckowstroemiidae are phoretic on passalid beetles in the New World. The family includes 37 species distributed in 4 genera, *Klinckowstroemia* (28 spp.), *Similantennurella* (2 spp.), *Klinckowstroemiella* (5 spp.) y *Antennurella* (2 spp.). Little is known about the true distribution and species diversity of these genera, especially the latter 3. We examined specimens of *Antennurella*, *Similantennurella* and *Klinckowstroemiella* collected from passalids from several entomological collections. We added Ecuador, French Guyana, Guatemala and Mexico to the distribution range of *Similantennurella*, Belize, Bolivia, Ecuador, Guatemala, Trinidad and Venezuela for *Antennurella*, and Bolivia, Ecuador, Honduras and Peru for *Klinckowstroemiella*. We report 8 new passalid host species previously unknown for these genera. Our study suggests a broad distribution range of the 3 genera, probably encompassing Mesoamerica and South America except Chile and Patagonia, where passalids are not found. *Antennurella*, *Similantennurella* and *Klinckowstroemiella* apparently occur primarily below 1500m altitude, while *Klinckowstroemia* has been repeatedly found above this limit. All genera have been found mostly on large passalids (>22 mm, long). We also cite the family for the first time from El Salvador.

**Keywords** Antennurella; Similantennurella; Klinckowstroemiella; Klinckoestroemia

### Introduction

Passalidae Leach 1815 is a family of mainly tropical xylophagic beetles (Schuster, 1978). Passalids spend virtually their entire life cycle living in tunnels inside decomposing logs (Villegas-Guzmán et al. 2012). They commonly form monogamous pairs and exhibit subsocial behavior and complex acoustic communication systems (Schuster & Schuster, 1985).

Currently, more than 200 species of mites have been reported to be associated with passalid beetles. They are distributed in 68 genera and 21 families in the Mesostigmata, Prostigmata, Astigmata and Oribatida suborders. Mites in 19 of these families, including Klinckowstroemiidae Camin & Gorirossi 1955 are known to form phoretic associations with their passalid host (Villegas-Guzmán et al. 2008). Klinckowstroemiid mites are known only from the New World and only from passalids but little else is currently known about the ecology and life cycle of its members.

The family Klinckowstroemiidae includes 37 species of mites, all associated with passalid beetles (Villegas-Guzmán et al. 2011) from Mexico to South America. Previous studies have shown an absence of host specificity by the mites in these phoretic associations (Villegas-Guzmán et al. 2012). The Klinckowstroemiidae includes 4 genera: *Similantennurella Rosario*

Received 26 August 2021

Accepted 01 July 2022

Published 22 May 2023

Corresponding author
Daniel Gygax<sup>D</sup>: dgygax90@gmail.com

Academic editor
Roy, Lise

https://doi.org/10.24349/1zn2-hq18

ISSN 0044-586X (print)

ISSN 2107-7207 (electronic)

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Schuster J. et al.

Licensed under
Creative Commons CC-BY 4.0

OPEN ACCESS

<footnote><sup>a</sup>Departamento de Biología, Universidad del Valle de Guatemala, 18 Avenida 11-95, 01015, Guatemala.</footnote>

<footnote><sup>b</sup>Programa Centroamericano de Maestría en Entomología, Universidad de Panamá.</footnote>
1988 (2 species), *Klinckowstroemiella* Türk, 1951 (5 species), *Klinckowstroemia* Baker & Wharton 1952 (28 species) and *Antennurella* Berlese 1903 (2 species). *Klinckowstroemia* (28 species) is widely distributed in Mesoamerica.

*Similantenurella spinata* Rosario 1988 was previously known from specimens collected in Costa Rica and Panama; *S. aspinata* Rosario 1988 is only known from the holotype collected in Costa Rica (Rosario, 1988). *Klinckowstroemiella* includes: *K. prima* Türk 1951, *K. helleri* Oudemans 1929, *K. blumae* Rosario & Hunter 1987, *K. sexisetosa* Rosario & Hunter 1987 and *K. trinidadis* Wisniewski & Hirschmann 1994 and is known from Brazil, Costa Rica, Panama, Surinam and Venezuela (Rosario & Hunter 1987). *Antennurella* includes two species, possibly synonyms: *A. trouessarti* Berlese 1904 and *A. tragardhi* Baker & Wharton 1952 (Rosario, 1988). However, the distribution range and passalid host spectra of this family is suspected to be much broader than previously reported.

The study aims to improve knowledge of the distribution and host spectra of klinckow-
stroemiid mites. We report new localities and host species for *Similantenurella*, *Klinckow-
stroemiella* and *Antennurella*. and we cite the presence of two species of *Klinckowstroemia* in
El Salvador, the first records of the family from this country.

**Material and methods**

We checked available literature to construct a database including all currently known countries and passalid host species in which the studied genera occur. We then examined mite species from the Entomology Collection of the Academy of Natural Sciences of Drexel University, Allan Gillogly personal collection, the Arthropod Collection of the Universidad del Valle de Guatemala, William F. Barr Entomological Museum at Idaho College, Florida State Collection of Arthropods, California Academy of Sciences Entomology Collection, as well as from passalids we collected which are deposited in the Arthropod Collection of the Universidad del Valle de Guatemala.

Passalids were also sampled in the Montecristo Trinational Protected Area (MTPA) located in northeastern Santa Ana Province, El Salvador (14° 25' N, y 89°23' O) between September and December 2015. This is the region where the borders of Guatemala, Honduras and El Salvador unite. The elevation of the region varies between 420 and 2418m. Passalids were collected by hand from decomposing logs in three altitudinal strata: cloud forest, mixed pine forest and tropical dry forest.

Beetles from each trunk were placed in vials with ethyl alcohol 70% labeled and taken to the Plant Protection Laboratory of the Faculty of Agronomic Sciences of the University of El Salvador. Later we removed the mites from the beetles and identified them to species using the key of Villegas-Guzmán et al. (2011). These specimens were deposited in the Entomology Collection of the University of El Salvador. Mites still attached to the host were removed with either fine tweezers or a fine brush. Mites were deposited in a smaller separate vial with ethyl alcohol 70%. For observation, mites were temporarily laid on a Petri dish or a microscope slide.

The diagnosis of genera is a summary based on Rosario (1988). *Antenurella*: Hyaline hood surrounding idiosoma; posterior portion of dorsal shield with 3 pronounced protuberances (Figures 1, 2, 3). *Similantenurella*: Hyaline hood forming crescent over gnathosoma and extending to or beyond level of anterior margin of anal shield, never surrounding entire idiosoma; posterior portion of dorsal shield with 3 protuberances, central protuberance much more pronounced than the other two (Figures 4, 5, 6). *Klinckostroemiella*: Hyaline hood less visible, forming a small cap anteriorly; anal shield fused to ventral shield; posterior part of shield without a protuberance (Figure 7). For identification of *Klinckowstreamia* specimens chaetotaxy was based on a taxonomic key published by Villegas-Guzmán et al. (2011).
<figure><img src="image_8.png" /><figcaption>**Figure 1** Close view of diagnostic protuberances of possible *Similantennurella* specimen (Photo by JCS).</figcaption></figure>

## Results

We collected 13 species of Passalidae in the MTPA, 6 species in cloud forest, four species in mixed pine forest and five species in the dry forest. Of 116 passalid individuals 21 had klinckowstroemiids (Table 1). Interestingly, klinckowstroemiid mites were found only in passalids collected from the mixed pine forest part of the MTPA, between 1059 and 2161 meters above sea level (14°24'9.72"N 89°22'52.82"O y 14°23'2.41"N 89°23'37.23"O).

### Klinckowstroemia atramaculata Rosario & Hunter, 1988

This species was associated with 15 individuals of *Heliscus eclipticus* Truqui and 4 of *Oileus sargi* Kaup collected in rotting trunks of *Pinus* and *Quercus*. Until now it has been reported only from Puntarenas, Costa Rica. *Klinckowstroemia atramaculata* presents the mesogynal shield (MgS) anteriorly rounded (Figure 4A); sternal seta 2 is smooth and sternal seta 1 is barbed. (Villegas-Guzmán *et al.*, 2011).

### Klinckowstroemia victoriae Rosario & Hunter, 1988

This species we found associated with two individuals of *Arrox agassizi* Kaup collected in trunks of *Quercus* sp. It has been reported from Xicotepec de Juárez (Puebla, México), Quetzaltenango Dept. y Puerta Parada in Guatemala. *Klinckowstroemia victoriae* presents sternal seta 4 short sternal seta 3 of medium length, one pair of setae a and sternal seta 1 is serrate latigynal shields united mesally with a wavy border (Figure 4B) (Villegas-Guzmán *et al.*, 2011).

<table><caption>**Table 1** Species of Klinckowstroemiidae and associated Passalid host collected in the MTPA in this study.</caption><thead><tr><th>Host species</th><th><em>Klinckowstroemia</em> species</th><th>females</th><th>males</th></tr></thead><tbody><tr><td><em>Heliscus eclipticus</em> (8)</td><td><em>K. atramaculata</em></td><td>26</td><td>31</td></tr><tr><td><em>Oileus sargi</em> (2)</td><td><em>K. atramaculata</em></td><td>24</td><td>26</td></tr><tr><td><em>Arrox agassizi</em> (1)</td><td><em>K. victoriae</em></td><td>6</td><td>7</td></tr></tbody></table>

The number in brackets after the passalid species is to the number of individuals of this species found with associated klinckowstroemiid mites
<figure><img src="image_9.png" /><figcaption>**Figure 2** Dorsal shield of *Antennurella* specimen. Note the posterior portion of dorsal shield with 3 pronounced protuberances. (Photo by K. Shnepp).</figcaption></figure>

al., 2011). *K. victoriae* is also the most common klinckwostroemiid species in some areas of the city and is associated to several passalid hosts (Table 2) (obs. pers.)

Several new localities and hosts were found for *Similantennurella*, *Antennurella*, *Klinckowstroemiella*. Five of the *Similantennurella* specimens examined (Table 3) were collected in previously unreported localities, including Ecuador, Mexico, French Guyana and Guatemala. All were found on *Passalus punctatiger* Lepeletier & Serville. Six of the *Antennurella* specimens examined (Table 3) belong to 6 new localities: Bolivia, Belize, Ecuador, Venezuela, Trinidad and Guatemala. Two males were found on *P. punctatostriatus* and a female was found on

<table><caption>**Table 2** Passalid hosts reported for *K. victoriae* in Guatemala.</caption><thead><tr><td>Host</td><td>Locality</td><td>Date</td><td>Collector</td></tr></thead><tbody><tr><td>Petrejoides guatemalae</td><td>Huehuetenango, 15km south of San Mateo Ixtatan, 2800m</td><td>6/V/1977</td><td>J. Schuster</td></tr><tr><td>Chondrocephalus granulifrons</td><td>Solola, Volcán Atitlán, 2500m</td><td>27/I/1977</td><td>C. McVean</td></tr><tr><td>Passalus punctatostriatus</td><td>Guatemala, Puerta Parada, 1840m</td><td>24/V/2018</td><td>J. Schuster</td></tr><tr><td>Arrox agassizi</td><td>Chiquimula, Trifiño, El Duraznal, 1668m</td><td>11/VI/2011</td><td>E. Cano</td></tr><tr><td>Verres hageni</td><td>Chimaltenango, Volcán Acatenango, 2000m</td><td>20/IX/1986</td><td>?</td></tr><tr><td>Ogyges laevssimus</td><td>Quetzaltenango, Fuentes Georginas, 2500m</td><td>5/VI/2015</td><td>E. Fuller</td></tr></tbody></table>
<figure><img src="image_10.png" /><figcaption>**Figure 3** *Klinckoswstroemiellaventral male: A – Region lacking abdominal suture; B – Hyaline hood not surrounding posterior body.*</figcaption></figure>

<figure><img src="image_11.png" /><figcaption>**Figure 4** *Antennurella* ventral male: A – Suture separating abdomen from rest of the body; B – Hyaline hood surrounding whole body. (Photos by JCS).</figcaption></figure>

*Verres furcilabris*, both previously unreported hosts (Table 3). Four of the *Klinckowstroemiella* specimens (Table 3) examined were from previously unreported localities: Bolivia, Peru, Honduras and Ecuador. Some specimens were collected from previously unreported hosts for the genus (Table 4).

**Discussion**

Of the Passalid beetles examined, approximately 180 were pinned individuals of which only 8 yielded Klinckowstroemiidae. Many mites fall off after submerging the host in alcohol or in collecting jars so this probably does not reflect true prevalence of the mites on these hosts. Klinckowstroemiids were found mostly on large passalid species such as *P. punctiger*
<figure><img src="image_12.png" /><figcaption>**Figure 5** Dorsal shield of 3 possible *Similantennurella* specimens. The top photo may be *Antennurella*. Note the wide hyaline hood that completely surrounds the body characteristic of *Antennurella*. Note the 3 protuberances, with the central protuberance much more pronounced than the other two as in *Similantennurella*. Also note the differences in form, size and position of the lateral protuberances between specimens. (Left and top photos by K. Shnepp, other by JCS) (Size line at lower right not valid).</figcaption></figure>

(30-35 mm), *P. interruptus* (35-45 mm) and *Verres* spp. (30-50 mm) (Table 4). The smallest hosts reported are *P. morio* (25-30 mm) and *P. punctatostriatus* (22-30 mm) as well as unidentified species of *Passalus* (*Pertinax*) (20mm). No *Klinckowstroemiella*, *Antennurella* or *Semilantennurella* specimens have been found on small passalids (less than 20 mm in length). Our study shows a broader distribution range for these three genera than previously known.

<table><caption>**Table 3** *Similantennurella*, *Antennurella*, *Klinckowstroemiella*, specimens with new locality/host records.</caption><thead><tr><th>Genus</th><th>Species</th><th>Host</th><th>Sex</th><th>Locality</th><th>Elevation</th><th>Date collected</th><th>Collector</th></tr></thead><tbody><tr><td>Similantennurella</td><td>S. spinata</td><td>P. punctiger</td><td>male</td><td>Esmeraldas, San Mateo, Ecuador</td><td>32m</td><td>19 XI 1956</td><td>F. H. Walz</td></tr><tr><td>Similantennurella</td><td>S. spinata</td><td>P. punctiger</td><td>male</td><td>Palenque, Chiapas, Mexico</td><td>61m</td><td>17 VII 1992</td><td>Mornburg</td></tr><tr><td>Similantennurella</td><td>S. spinata</td><td>P. punctiger</td><td>2 males</td><td>Acayucan, Veracruz, Mexico</td><td>102m</td><td>13 X 1957</td><td>B. y K. Dreisbach</td></tr><tr><td>Similantennurella</td><td>S. spinata</td><td>P. punctiger</td><td>male, female</td><td>Ixpanpajul, Peten, Guatemala</td><td>125m</td><td>28 V 2017</td><td>E. Fuller</td></tr><tr><td>Similantennurella</td><td>S. spinata</td><td>P. punctiger</td><td>male</td><td>Amazonas Nature Lodge, French Guyana</td><td>300m</td><td>04 I 2016</td><td>J. Eger, J. Wappes, R. Morrris</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. punctatostriatus</td><td>2 males</td><td>8km south of Montepío, Veracruz, Mexico</td><td>150 m</td><td>9 X 1997</td><td>Gillogly</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. punctiger</td><td>female</td><td>Toledo, Columbia Forest Reserve, Belize</td><td>below 825m</td><td>8 VIII 2006</td><td>P.W. Kovarik</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. interruptus</td><td>male</td><td>Aragua, Maracay, La Cumbre, Venezuela</td><td>1200m</td><td>5 VIII 1992</td><td>unknown</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. punctiger</td><td>female</td><td>Quetzaltenango, Volcán St. María, Guatemala</td><td>1250m</td><td>23 II 1977</td><td>unknown</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. punctiger</td><td>female</td><td>Mochiflan, Acahuizotla, Guerrero, Mexico</td><td>650m</td><td>08 IV 1996</td><td>L. Delgado</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>V. furcilabris</td><td>female</td><td>George County Mountains, Trinidad</td><td>below 940</td><td>24 III 1987</td><td>unknown</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. punctiger</td><td>2 males</td><td>47km South of Santo Domingo, Los Ríos, Ecuador</td><td>220m</td><td>26 XIII 1997</td><td>A. Gillogly</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. punctiger</td><td>male, female</td><td>Estación Biológica las Guacamayas, Petén, Guatemala</td><td>70m</td><td>28 VI 2014</td><td>J. Schuster</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. interruptus</td><td>female</td><td>Santa Cruz, Bolivia</td><td>500m</td><td>16-21 X 2006</td><td>unknown</td></tr><tr><td>Antennurella</td><td>A. trouessarti</td><td>P. (Pertinax) sp.</td><td>unknown</td><td>San Luis-Canchacan, Petén, Guatemala</td><td>300m</td><td>19-VII-2008</td><td>C.W. & L.B. O'Brien & F. Skillman</td></tr><tr><td>Klinckowstroemiella</td><td>K. sp</td><td>P. convexus</td><td>female</td><td>Limoncocha, Ecuador</td><td>215m</td><td>10 VIII 1997</td><td>A. Gillogly</td></tr><tr><td>Klinckowstroemiella</td><td>K. blumae</td><td>P. punctiger</td><td>3 females</td><td>Blues Lagoon, Honduras</td><td>?</td><td>25 IV 1947</td><td>C.W. Cook</td></tr><tr><td>Klinckowstroemiella</td><td>K. helleri</td><td>P. interruptus</td><td>male</td><td>Rio Monzon, Huanuco, Peru</td><td>778m</td><td>21 II 1970</td><td>J. Schuster</td></tr><tr><td>Klinckowstroemiella</td><td>K. sp.</td><td>V. sinuaticollis</td><td>male</td><td>Veraguas, Panama</td><td>850m</td><td>11 X 1992</td><td>A. Gillogly</td></tr><tr><td>Klinckowstroemiella</td><td>K. prima</td><td>P. morio</td><td>male</td><td>Sucumbios, Ecuador</td><td>215m</td><td>12 VIII 1997</td><td>A. Gillogly</td></tr><tr><td>Klinckowstroemiella</td><td>K. sexisetosa</td><td>V. aspina</td><td>female</td><td>Los Ríos, Rio Palenque, Ecuador</td><td>below 170m</td><td>18 VI 1974</td><td>J. Longino</td></tr><tr><td>Klinckowstroemiella</td><td>K. sexisetosa</td><td>V. longicornis</td><td>male, female</td><td>Cartago, Turrialba, Costa Rica</td><td>600m</td><td>12 VII 1965</td><td>Allsop</td></tr><tr><td>Klinckowstroemiella</td><td>K. blumae</td><td>P. interruptus</td><td>female</td><td>Santa Cruz, Bolivia</td><td>500m</td><td>16-21 X 2006</td><td>unknown</td></tr></tbody></table>
<figure><img src="image_13.png" /><figcaption>**Figure 6** Dorsal shield of *Klinckowstroemiella*. Note the posterior part of shield without a protuberance and the presence of a deep marginal groove around the dorsal shield latero posteriorly (arrow) (Photo by K. Shnepp).</figcaption></figure>

Our findings suggest a distribution encompassing Mesoamerica and South America except Chile and Patagonia, where passalids are not found. Furthermore, all the localities of the specimens of three of these genera (*Klinckowstroemiella*, *Antennurella* or *Semilantennurella*) examined were found below 1500 m above sea level. In contrast, *Klinckowstroemia* has been repeatedly found above this limit.

As a final note, the diagnosis of genera is based on Rosario (1988), including the posterior portion of dorsal shield with 3 protuberances with the central protuberance much more pronounced than the other two as diagnostic feature of *Similantennurella*. Figure 4 shows this feature in 3 different klinckowstroemiids. However, a marked variation in shape, size and position of the protuberances is present, suggesting 3 different (possibly undescribed) species.
<figure><img src="image_14.png" /><figcaption>**Figure 7** Close view of diagnostic protuberances of *Antennurella* (Photo by JCS).</figcaption></figure>

<table><caption>**Table 4** Passalid host species reported for *Antennurella*, *Similantennurella* and *Klinckowstroemiella*.</caption><thead><tr><th>Antennurella</th><th>Similantennurella</th><th>Klinckowstroemiella</th></tr></thead><tbody><tr><td><strong>Passalus coniferus</strong> Eschscholtz</td><td rowspan="5"><strong>P. punctiger</strong></td><td><strong>P. convexus</strong> Dalman</td></tr><tr><td><strong>P. interruptus</strong> Linnaeus</td><td><strong>P. interruptus</strong></td></tr><tr><td><strong>P. punctatissimus</strong> Eschscholtz</td><td><strong>P. jansoni</strong> Bates</td></tr><tr><td><strong>P. punctatostriatus</strong> Percheron</td><td><strong>P. morio</strong> Percheron</td></tr><tr><td><strong>P. punctiger</strong> Lepeletier &amp; Serville.</td><td><strong>P. (Pertinax)</strong> sp. Kaup</td></tr><tr><td><strong>P. rusticus</strong> Percheron</td><td rowspan="2"><strong>Verres longicornis</strong> Luederwaldt</td></tr><tr><td><strong>Popilus</strong> sp. Kaup</td></tr><tr><td><strong>Ptichopus angulatus</strong> Percheron</td><td><strong>V. hageni</strong> Kaup</td></tr><tr><td><strong>Verres furcilabris</strong> Eschscholtz</td><td><strong>Veturius aspina</strong> Kuwert</td></tr><tr><td></td><td><strong>V. platyrhinus</strong> Westwood (V. sp)*</td></tr><tr><td></td><td><strong>V. sinuaticollis</strong> Kuwert</td></tr></tbody></table>

Hosts reported in this study for the first time are showed in bold letters: * The V. platyrhinus specimen (as reported by Rosario and Hunter 1987) was found in Costa Rica, however this species is endemic to Colombia (Boucher, 2005).

## Acknowledgements

JCS would like to thank the late Preston Hunter for the stimulus to study these mites and for some species determinations and literature. We thank Barry OConnor for literature and access to the University of Michigan collection. Also, to Alan Gillogly for access to his passalid collection and to Jon Gelhaus and the Philadelphia Academy of Sciences and the Universidad del Valle de Guatemala for access to their respective passalid collections. We also thank various students and other people who have brought us passalid specimens or helped in mite studies, and especially thanks to Ruben Morales, Ed Fuller, Elinor Stevens, Paul Skelley and Kyle Schepp for collecting passalids, mites and/or photos of mites. We appreciate Jerry Krantz, Barry OConnor, Margarita Palmieri and Paula Muñoz for reviewing the manuscript.
ORCID

Jack Schuster https://orcid.org/0000-0001-5895-7639

Daniel Gygax https://orcid.org/0009-0005-2362-0063

Francisco Serrano-Peraza https://orcid.org/0000-0002-1467-1070

References

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