Research Article |
Corresponding author: Frédéric Beaulieu ( frederic.beaulieu@agr.gc.ca ) Corresponding author: Edwin Javier Quintero-Gutiérrez ( ejquinterog@gmail.com ) Academic editor: Farid Faraji
© 2019 Frédéric Beaulieu, Edwin Javier Quintero-Gutiérrez, Dorotee Sandmann, Bernhard Klarner, Rahayu Widyastuti, Orlando Cómbita-Heredia, Stefan Scheu.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Beaulieu F, Quintero-Gutiérrez EJ, Sandmann D, Klarner B, Widyastuti R, Cómbita-Heredia O, Scheu S (2019) Review of the mite genus Ololaelaps (Acari, Laelapidae) and redescription of O. formidabilis Berlese. ZooKeys 853: 1-36. https://doi.org/10.3897/zookeys.853.29407
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A species of laelapid mite, Ololaelaps formidabilis, is redescribed based on male and female adults from soil in Sumatra, Indonesia. This species is distinguished from other Ololaelaps species by its metapodal platelet narrowly fused with the parapodal plate and by its hologastric shield having two inverted-V-like ridges. The genus is redescribed based on a review of the literature and examination of specimens of some species. Valid species of Ololaelaps are listed and accompanied by notes on morphological characters to assist future revision of the genus.
Gamasida, Indonesia, laelapid mites, rubber plantation, soil fauna, Sumatra
Laelapidae is a large, ecologically diverse family of Mesostigmata, with several species described from Indonesia, including symbionts of bees (
The cosmopolitan genus Ololaelaps was initially proposed by
The initial goal of this paper was to redescribe O. formidabilis, which was collected from soil in a rubber plantation (Hevea brasiliensis Müll. Arg.) near a lowland rainforest on the island of Sumatra, Indonesia. We took this opportunity to review the generic concept, based on the literature and examination of specimens of some species. We also present (1) a list of valid species of Ololaelaps, including notes on their most salient morphological features, and (2) a list of species that were previously classified as Ololaelaps but that have unclear taxonomic affinity (nomina dubia) or that now belong to other genera of Mesostigmata.
List of valid Ololaelaps species (in bold) and synonyms (in regular font), type localities [and additional records], habitats and depository.
Species | Original genus | Type locality [other distribution records] | Type habitat [other records] | Type repository1 | Notes and additional references (incl. for selected distribution records) |
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bregetovae Shereef & Soliman, 1980: 81 | Ololaelaps | EGYPT: Giza | debris |
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burdwanensis Bhattacharyya, 1978: 86 | Ololaelaps | INDIA: Burdwan (West Bengal) | soil under grass beside pond |
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caucasicus Bregetova & Koroleva, 1964: 73 | Ololaelaps | RUSSIA: near Kizlyar (Dagestan); ARMENIA: Yerevan2 | litter of Elaeagnus (Russia), litter under ash tree (Armenia) |
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confinis Berlese, 1904: 261 [?syn. of placentula] | Ololaelaps | NORWAY | ? |
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Not illustr. by |
dililoensis Marais & Loots, 1972: 31 | Ololaelaps | REPUBLIC OF THE CONGO: Eala | soil |
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expansus Ma, 2015: 95 | Pristolaelaps | TAIWAN: Tainan | soil |
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flavus Ewing, 1909: 66 [syn. of placidus] | Laelaps | USA: Arcola (Illinois) | under log |
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Lectotype designated by |
formidabilis Berlese, 1913: 82 | Ololaelaps | INDONESIA: Semarang (Java); [Sumatra (this paper)] | ? [forest litter] |
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gamagarensis Jordaan & Loots, 1987: 49 [syn. of mooiensis] | Ololaelaps | SOUTH AFRICA: Gamagara River, Sishen | soil under grasses and reeds, on river bank |
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Synonymy by |
haemisphaericus Koch, 1839b: 16 [?syn. of sellnicki] | Iphis | GERMANY | marshy meadows | ? | Recognized as Ololaelaps by |
halaskovae Bregetova & Koroleva, 1964: 81 [syn. of venetus] | Ololaelaps | RUSSIA (widespread locations); UKRAINE (Zakarpattia Oblast); MOLDAVIA: Egorovka2 | litter in meadows and forests; on small rodents or in their nests |
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Synonymy by |
hemisphaera Berlese, 1916b: 303 | Ololaelaps | USA: Columbia (Missouri4) | litter |
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holaspis Oudemans, 1902b: 53 | Hypoaspis | ITALY: Sanremo | litter |
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interruptus Karg, 1994: 186 | Pseudoparasitus | ECUADOR (Galápagos Islands): Cerro Banderas, 4 km NE of Santa Rosa, Santa Cruz island | litter of Miconia sp., in a cave |
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leptochelae Karg, 1994: 187 | Pseudoparasitus | ECUADOR (Galápagos Islands): near El Puntudo, Santa Cruz island | moist litter in fern-sedge zone |
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magnichelas Ewing, 1909: 65 [syn. of placidus] | Laelaps | USA: Muncie (Illinois) | moss |
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Lectotype designated by |
mooiensis Ryke, 1962: 126 | Ololaelaps | SOUTH AFRICA: Mooi River, Potchefstroom; [ANGOLA, IRAN] | damp soil on river bank; [soil, litter] |
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nasri Hassan, 1989: 593 | Ololaelaps | EGYPT: Kafr Shokr | debris under citrus trees | ? | |
obovatus Womersley, 1960: 33 | Pristolaelaps | AUSTRALIA: Koroit (Victoria) | ? | SAM | |
paratasmanicus Ryke, 1962: 127 | Ololaelaps | NEW ZEALAND: Dunedin; [CHINA: Kunming] | bracken |
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placentula Berlese, 1887: 3 | Laelaps | ITALY: Vallombrosa; [widespread in Europe; CHINA, RUSSIA, USA, CANADA] | moss; [litter in forests and meadows, nest of small mammals] |
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placidus Banks, 1895: 128 | Laelaps | USA: near Roslyn (New York); [CANADA] | wet moss; [litter] |
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Lectotype designated by |
platensis Berlese, 1916a: 166 | Ololaelaps | ARGENTINA: La Plata | ? |
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rectagoni Karg, 1993b: 269 | Pseudoparasitus (Ololaelaps) | ECUADOR (Galápagos Islands): south of Wreck Bay, San Cristóbal island | moist and salty litter |
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sellnicki Bregetova & Koroleva, 1964: 77 | Ololaelaps | RUSSIA, UKRAINE, LITHUANIA2; [widespread in western parts of Eurasia] | wet meadows, stream banks, coastal habitats, alpine meadows, rodent nests |
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sinensis Berlese, 1923: 252 | Ololaelaps | CHINA: near Beijing | ? |
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Originally described as O. venetus var. sinensis; |
sitalaensis Bhattacharyya, 1978: 84 | Ololaelaps | INDIA: Sonarpur (West Bengal) | litter under Pistia stratiotes at pond margin |
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tasmanicus Womersely, 1956: 571 | Pristolaelaps | AUSTRALIA: Tasmania; [USA: Hawaii; NEW ZEALAND] | strawberry plants; [moss, soil, on a rat] | SAM |
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translineatus Barilo, 1991: 15 | Pseudoparasitus (Ololaelaps) | UZBEKISTAN: Baysun | turf of [urban] park |
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ussuriensis Bregetova & Koroleva, 1964: 75 | Ololaelaps | RUSSIA (Primorsky Territory)2; [CHINA] | on small rodents, in their nests, or soil |
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venetus Berlese, 1903: 14 [?jun. syn. of placidus] | Laelaps (Hypoaspis) | ITALY: Veneto3; [widespread in Europe and parts of Asia] | moss; [see records for O. halaskovae] |
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Laelaps (H.) venetus was proposed by |
wangi Bai, Gu & Wang, 1996: 74 | Ololaelaps | CHINA: Southern Yinchuan; [SOUTH KOREA] | decaying Zea mays; [grassland soil] | EDC |
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This study is part of a larger investigation on arthropods of Indonesia within the framework of the interdisciplinary project “Ecological and socioeconomic functions of tropical lowland rainforest transformation systems (Sumatra, Indonesia)” – EFForTS. For details on the study region and the experimental design, see
Soil and litter samples were taken, using a spade, from rubber plantation plots at the rainforests of Bukit Duabelas (National Park) and Harapan (National Forest), Jambi Province, Sumatra (see “Material examined” section for details on localities). Samples represented 16 × 16 cm of litter and soil taken down to a 5-cm depth. Mites were extracted from samples using a modified high-gradient canister method (
Photographs and measurements were made using a compound microscope (Nikon Eclipse Ci or Leica DM5500B) equipped with phase contrast or differential interference contrast and connected to a computer-controlled digital camera (Sight Ds-L3 or Leica DMC4500). Most images were captured in stacks (with focal depth manually or electronically controlled). Selected images were combined using Zerene Stacker version 1.04 or Helicon Focus 6.7.1 Pro (Helicon Soft Ltd., 2000). Digital drawings were prepared using Adobe Illustrator, version CC 2015 (19.0.0), based on mite photographs that were first imported into the software.
All measurements are given in micrometers (μm) and presented as ranges (minimum–maximum). Lengths of shields were measured along their midlines, and widths at the widest point except for the sternal shield, measured at level of setae st2. Legs were measured from proximal margin of the coxa to the tip of tarsus, excluding ambulacrum (stalk, claws, pulvillus), and corniculi from their apex to the midpoint of their internal base. Spermatodactyl was measured from its point of departure from the movable digit to its apex. Notations of structures and idiosomal chaetotaxy generally follow
Specimens of O. formidabilis are deposited in LIPI (Indonesian Institute of Science), Cibinong, Indonesia; the
Additional photos of the species are digitally deposited in the online database available at ecotaxonomy.org.
The diagnosis and description of the genus were prepared after consultation of previous diagnoses of the genus (
Pristolaelaps
Womersley, 1956: 571. Synonymy by
Laelaps (Hypoaspis) venetus Berlese, 1903
(adult male and female, unless stated).Well-sclerotized hypoaspidine laelapid with a hologastric (genitiventrianal) shield in female, bearing 3–5 pairs of preanal setae (plus st5), as well as the following character states: dorsal shield covering entirely idiosoma dorsally, narrowly to broadly extending onto venter; bearing 39 or slightly fewer pairs of slender setae, including px2–3 and often one Jx. A pair of well-sclerotized presternal platelets. Female with seta st4 on sternal shield or on soft cuticle (or putatively on endopodal plate). Peritrematal shield free posteriorly or variously (narrowly) fused with hologastric and/or parapodal shields, via metapodal platelet; metapodal platelet free or variously fused to above-mentioned shields; parapodal plate well-developed, subtriangular. Soft opisthogastric cuticle with 5–10 pairs of setae. Male holoventral shield broad, fused to parapodal-exopodal plates, sometimes also to peritrematal shield. Gnathotectum convex, with few to numerous fine denticles; deutosternal groove with six rows of 1–10 denticles; female cheliceral movable digit with two teeth (rarely more), fixed digit with 3–5 (exceptionally 8); palp-apotele three-tined, third tine reduced. Leg chaetotaxy normal for Laelapidae; setae generally slender.
Dorsal idiosoma.
Dorsal shield relatively large (435–800 in female), broadly oval to narrowly suboval (length/width ratio 1.2–1.8), completely covering idiosoma dorsally, barely to moderately extending ventrally (this can be determined most accurately before slide-mounting); shield smooth (except for fine granulation or punctuation) to strongly reticulate; shield’s ventral extension (‘epipleura’ of
Ventral idiosoma.
Tritosternum normal, with two pilose laciniae. Presternal region with a pair of sclerotized platelets, wedge-shaped to subrectangular, lineate (typically with 2–4 transversal lineae); typically an additional, poorly sclerotized area, lineate and granulate, anteriorly or anteromesally adjoining each platelet. Female sternal shield as long as or longer than wide, sometimes wider than long; shield length/width ratio 0.6–1.8; Shield posterior margin straight, slightly to moderately concave, or sometimes convex; shield reticulate, smooth in its posterior fourth or fifth, bearing 3–4 pairs of simple setae and 2–3 pairs of poroids, therefore sometimes including seta st4 and poroid iv3; seta st4 on sternal shield (near or on its posterolateral edge), on soft cuticle, or apparently on endopodal plate (Table
Gnathosoma.
Gnathotectum with subtriangular to rounded margin, usually finely denticulate, may appear smooth when denticles sparse or (possibly) absent. Deutosternal groove of moderate, regular width, or slightly tapering posteriorly, with six (occasionally seven, and rarely five) rows of denticles, each row bearing 1–10 denticles, most of the rows with 3–7 denticles; denticulate rows usually preceded by a smooth ridge anteriorly, and sometimes also posteriorly. Corniculi horn-like, of moderate length. Internal malae with two pairs of long projections, median pair fimbriate on its basal portion, lateral pair smooth or branched or fimbriate in its apical portion; lateral projection absent in males (and apparently in the female of O. sitalaensis). Palptarsal claw three-tined, third tine reduced. Chelicerae of moderate length, chelate-dentate; female movable digit with two moderately-sized teeth, rarely more (two additional small teeth between the two typical large teeth in O. interruptus; Table
Legs.
Chaetotaxy normal for Laelapidae (sensu
Spermatheca.
Spermathecal ducts well-sclerotized and discernable in some species.
Some diagnostic features of valid Ololaelaps species based on the literature, except for a few species for which type (**) or voucher (*) specimens were examined. Species are sorted in groups based on shared features, mainly the various fusion of shields ventrally (groups may or may not reflect relatedness).
Species | Shared features (mostly fusion of shields1) | Dorsal shield ornamentation1 | Epipleura2 ornament. | Dorsal seta Jx | Insertion of st43 | Other features1 | Notes and references (redescriptions) |
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venetus | (1) all shields (HOLOG + METAP + PERIT + PARAP) narrowly fused together; (2) spermatod. with sinuous duct; (3) spermathecae well-sclerotized, distinctive | smooth with sculptured areas anteriorly (Evans and Till’s text) | smooth | 1 | stern. | JV3, ZV2 setae sometimes off HOLOG |
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placidus * | smooth except light reticul. near ant. margin | smooth | 0–1 | stern. | as above |
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sellnicki | as venetus; reticul. visible only when freshly moulted ( |
smooth?4 | 1 | stern. | JV3, ZV2 off HOLOG; z1, z3 absent |
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hemisphaera | HOLOG + METAP + PERIT fused [PARAP apparently free] | ? | lineate-reticulate? | ? | soft cut.? | broad idiosoma | not illustr. in |
interruptus | (1) HOLOG + METAP + PERIT narrowly fused [PARAP clearly free]; (2) PERIT notched post. | ? | ? | 1 | soft cut. | MD with 2 small teeth in-between the 2 standard teeth; broad idiosoma | |
leptochelae | ? | ? | ? | ? | FD with a total of 8 teeth | ||
burdwanensis | HOLOG + METAP + PARA narrowly fused [PERIT free] | ? | lineate-reticulate | 1 | soft cut.? | ||
translineatus | smooth? | lineate-reticulate | 1 | soft cut.? | sternal shield with transverse ridge; spermatod. at 90° angle from MD; spermath. distinctive | similar to O. burdwanensis | |
wangi | smooth except lineate anteriorly | lineate-reticulate | 0 | soft cut.? | only 2–4 deutosternal denticles / row | similar to O. burdwanensis; |
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formidabilis *,** | only METAP + PARAP fused | light reticul.; lighter and sparser anteriorly | reticulate | 0–1 | soft cuticle | HOLOG with inverse V-shaped ridges; spermatod. elongate; spermath. not discerned |
O. formidabilis sensu |
caucasicus | only HOLOG + METAP (partly to completely) fused | similar to placentula or ussuriensis? | lineate-reticulate | 0–1 | stern. or soft cut. | broad idiosoma; spermatheca not discerned | similar to O. ussuriensis; |
dililoensis | dense scale-like reticul. post., smooth or scattered reticul. ant. | reticulate | 0 | soft cut. | broad idiosoma | ||
holaspis | only HOLOG + METAP (partly to completely) fused | reticulate? (Oudemans’ text says “all shields with large scales”) | ? | ? | soft cut.? | elongate idiosoma |
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mooiensis | reticulate; reticul. sparser anteriorly | ? | 0–1 | soft cut. or endop.? | elongate idiosoma; METAP rarely free (based on syn. O gamagarensis) |
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placentula * | essentially smooth (finely granulate) or faintly reticulate | lineate-reticulate | 0 | stern. | broad idiosoma; sternal shield wider than long, with concave margin; PERIT reaching past coxa IV; spermatheca not discerned |
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platensis | ? | ? | ? | soft cut.? | peritreme short, reaching between coxae I–II; ZV1 absent? |
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rectagoni | ? | ? | 0 | soft cut.? | j1 seta elongate; broad idiosoma and HOLOG; PARAP truncate; ZV3 apparently on HOLOG |
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sinensis | ? | ? | ? | soft cut.? |
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ussuriensis | polygonal reticul. scarcely evident (text) | lineate-reticulate | 0? | stern. | spermatheca not discerned; only 2–3 deutosternal denticles / row |
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bregetovae | all shields (HOLOG, METAP, PERIT, PARAP) free | with (scale-like?) reticulation post. | ? | 0? | ? | elongate idiosoma | similar to O. tasmanicus and O. sitalensis? |
expansus | ? | ? | 0? | soft cut. | |||
nasri | finely granulate? | lineate-reticulate? | 0 | soft cut.? | broad dorsal and sternal shields | similar to O. obovatus | |
obovatus | smooth? | ? | ? | soft cut. | broad idiosoma; ZV1 absent? | ||
paratasmanicus | reticulate | ? | 0 | soft cut. | elongate idiosoma; HOLOG rounded laterally | similar to O. tasmanicus; |
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sitalaensis | ? | reticulate | 1 | soft cut.? | elongate idiosoma | ||
tasmanicus | lightly reticulate (Womersley’s text) | ? | 0 | soft cut. |
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We herein recognize 26 valid species names in the genus Ololaelaps, and at least four synonyms (Table
The identity of Iphis haemisphaericus (
List of species that have been previously considered in Ololaelaps (as genus or subgenus), but herein excluded or considered dubious species (nomina dubia).
Species | Original genus | Current genus | Key sources for current placement | Sources placing it in Ololaelaps | Additional notes | Type locality |
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coleoptratus Berlese, 1888: 198 | Hypoaspis | Hydrogamasellus (Ologamasidae) |
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ARGENTINA: Buenos Aires |
festivus Koch, 1839b: 8 | Zercon | nomen dubium | – |
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See also |
GERMANY: Neumarkt |
germanicus Karg, 1965: 277 | Ololaelaps (Cypholaelaps) | Pseudoparasitus (Laelapidae) |
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GERMANY: Zörbig | |
globulus Koch, 1839b: 17 | Iphis | nomen dubium | – |
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GERMANY: Regensburg |
haemisphaericus Koch, 1839b: 16 | Iphis | nomen dubium: either Stylochirus (Ologamasidae) or Ololaelaps (as syn. of O. sellnicki; see Table |
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GERMANY |
haemisphaericus Berlese, 1916a: 166 | Ololaelaps (Cypholaelaps) | nomen dubium; note that Cypholaelaps semiglobulus |
– |
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ARGENTINA: La Plata |
holostaspoides Canestrini, 1884: 700 | Laelaps | Sessiluncus (Ologamasidae) |
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Unclear if types have been examined, but |
ITALY: Messina |
inornatus Johnston, 1849: 305 | Eumaeus | nomen dubium | – | considered a sen. syn. of Ololaelaps confinis in |
See notes for Z. festivus above, and |
UK (Scotland): Berwickshire |
pergibbus
Berlese (in |
Ololaelaps? | species name not available | – |
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The species name was not published (see |
CHINA |
Meanwhile,
There is no indication that anyone examined Koch’s types of haemisphaericus, and the types of most species described by Koch are presumably lost. Therefore, it may be impossible to confirm with certainty whether Koch’s species is Stylochirus or Ololaelaps. Resolving this dual identity of Iphis haemisphaericus (
Some species names once considered as Ololaelaps are herein excluded from the genus, based on the interpretation of the original description or more recent publications (Table
Although
Dorsal shield broad, length/width ratio ~1.3–1.4, lightly reticulate, bearing 39 pairs of simple setae, including px2–3, plus one unpaired seta Jx (sometimes absent); all setae short (21–27; j1, z1, J5 shorter); shield with gland opening gd4 conspicuous, on shield margin; epipleura narrow, strongly reticulate. Female sternal shield as long as wide (length/width ratio 0.96–1.02), bearing setae st1–st3; seta st4 and poroid iv3 on soft cuticle. Hologastric shield with two inverted V-like ridges, and strongly reticulate; cells scale-like in region anterior to anus, bearing seta st5 and five pairs of preanal setae. Soft opisthogastric cuticle laterad of shield with nine pairs of setae. Peritrematal shield free posteriorly, reaching level of coxa IV posterior margin. Metapodal shield suboval, narrowly fused to parapodal shield (and contiguous with hologastric shield) in female. Deutosternal groove with 3–5 denticles per row. Spermatodactyl prominent, 1.8× as long as movable digit.
(Figs
Idiosomal venter
(Figs
Ololaelaps formidabilis, adult female. A ventrolateral region of idiosoma, showing the well-reticulated epipleuron (ventrolateral portion of dorsal shield), gland opening gd4, and the dorsal shield’s marginal strip (“strip”); note that the epipleuron appears broader than in live specimen, because the specimen was squashed on the slide, as indicated by the broken dorsal shield B central region of the dorsal shield, showing the light reticulation of the opisthonotal area (near J1, Jx) and even lighter reticulation of the podonotal area (see between setae j5) C–E metapodal platelet (arrow), variously fused to the parapodal plate and contiguous with the hologastric shield. Scale bars: 50 µm (A); 100 µm (B); 50 µm (C–E).
Gnathosoma
(Fig.
Legs
(Fig.
Spermatheca. Not discerned.
(Figs
Idiosomal venter
(Fig.
Gnathosoma
(Fig.
Legs. Chaetotaxy and setae thickness similar to that of female. Lengths of legs: I 406–415, II 301–310, III 295–305, IV 380–395.
INDONESIA, Sumatra • 1♀, Harapan rainforest, litter from rubber tree plantation, research site HR4b, 01°48'18"S, 103°15'52"E, 71 m a.s.l. (LIPI; internal project ID macrolitterHR4b13_MESOS1_1) • 1♀, same data as preceding (CNC1098357; internal project ID macrolitterHR4b13_MESOS1_2) • 1♀ (with an egg), Bukit Duabelas rainforest, litter in rubber tree plantation, research site BR4b, 02°04'36"S, 102°46'22"E, 51 m a.s.l. (
Our discovery of Ololaelaps formidabilis in Sumatra appears to be the second record of the species in Indonesia, the first corresponding to the original description by Berlese from Java specimens. It is unique among described species of Ololaelaps in having its metapodal platelet fused to the parapodal plate and free from the peritrematal and hologastric shields. Note, however, that the metapodal platelet is tightly contiguous with the hologastric shield and that in some specimens, at some focal depth, it may even appear narrowly fused with it (Fig.
The male holotype of O. formidabilis (
Ololaelaps formidabilis A male holotype (slide 145/29): ventral view, with arrow pointing at spermatodactyl B female paratype (slide 145/30): region of hologastric shield, showing two inverted V-shaped ridges (v), and the ‘bridge’ (br) connecting parapodal (par) and metapodal (met) plates. Photographs courtesy of Roberto Nannelli.
At present, Ololaelaps appears as a relatively well-defined genus, characterized by a unique combination of characters, many of which, individually, are shared with other genera of Laelapidae, especially hypoaspidines. The most unique feature of Ololaelaps is the female genital shield hyperdeveloped posteriorly and fused with the anal shield to occupy most of the opisthogaster and capture 3–5 pairs of setae in addition to st5 and circumanals. The genital shield is also expanded in several other genera (e.g., Laelaspis, Laelaspisella, Pseudoparasitus, Pogonolaelaps;
The hemispherical nature of the idiosoma of several species of Ololaelaps is also distinctive. However, this attribute may have led to misidentifications or misclassifications in the past, as some species in other families, especially Ologamasidae, have a similarly glossy, dome-shaped dorsal shield (see Table
As explained in
An additional set of features that further distinguish some Ololaelaps species from other laelapid genera is the various fusions of the peritrematal, parapodal, metapodal and hologastric shields. Even for groups with opisthogastric (i.e., genitiventral) shields such as Laelaspis and Pseudoparasitus, we are not aware of such fusion among shields. The peritrematal and parapodal shields, however, are coalesced in a few other laelapids, such as Nidilaelaps annectans (Womersley) (
The female of some Ololaelaps species have seta st4 and poroid iv3 on the sternal shield. This is rare in laelapids, although common within other groups, especially Rhodacaroidea. Seta st4 is also born on the sternal shield (complex) in groups where the shield is fused posteriorly with endopodals (e.g., many ologamasids and pachylaelapids). However, in Ololaelaps, this feature seems associated with the anterolateral expansion of the genital shield, which leaves little soft cuticle available for the insertion of st4 and iv3. In other genera where the genital shield is more pronounced anteriorly, st4 has even disappeared (
The males of Ololaelaps are not as distinctive as females, although they can be distinguished from those of most other laelapid genera by the degree of development of the holoventral shield posterolaterally and its fusion to parapodal-exopodal shields, and sometimes to the peritrematal shields. However, a similar ventral shield arrangement occurs in the males of other laelapids, for instance N. annectans (
While it may be easy to identify a given Ololaelaps mite to genus, it is more difficult to identify it to species. Examination of types, as well as a critical assessment of intraspecific variation based on additional specimens will be necessary to clarify species boundaries and uncover synonymies. In particular, the following characters should be scrutinized during species (re)descriptions.
The type of fusion between metapodal, peritrematal, parapodal, and hologastric plates appears as a useful starting point to initiate species identification, because it sorts species into broad groups, which are phylogenetically meaningful in some cases (Table
(1) Fused peritrematal-hologastric-parapodal shields by way of the metapodal platelet. Note that the parapodal plate ranges from clearly to ambiguously fused to, or merely tightly contiguous with, the metapodal ‘bridge’ (e.g., compare figs 74–76 in
(2) Spermathecal ducts (= tubuli annulati,
(3) Spermatodactyl with a sinuous duct, and a subapical hump or bend (
(4) Dorsal shield with narrow, smooth epipleura (i.e., ventrolateral extensions of the dorsal shield) vs broad, lineate-reticulate epipleura of the placentula group. Other species may have narrow epipleura, smooth or reticulate but descriptions are often lacking in such details, in part because determining the extent of the epipleura is most readily done before slide-mounting of the specimen (
(5) A fifth character associated with the venetus group is the insertion of setae JV3 and ZV2 off the hologastric shield in O. sellnicki and in some individuals of O. venetus and O. placidus (Table
The placentula group was defined by four characters (three mentioned by
(1) six other species have the metapodal platelet fused to the hologastric shield (and free from parapodal/peritrematal plates), making this type of fusion relatively common in the genus (Table
(2) a poorly sclerotized (i.e., inconspicuous) spermatheca may characterize other species, given that it has been described in five species only (see above);
(3) at least two other species have the spermatodactyl with a non-sinuous duct (see above); and
(4) several other species have reticulate or lineate-reticulate epipleura that at least superficially resemble those of the placentula group of species. The ventral extent of the epipleura and its exact type of ornamentation should be scrutinized for each species. Members of the placentula group, O. placentula, O. ussuriensis and presumably O. caucasicus (note that O. ussuriensis and O. caucasicus were not illustrated dorsally) have a dorsal shield smooth or faintly reticulate, in contrast to conspicuously lineate-reticulate epipleura, which are relatively well extended ventrally (Table
Ololaelaps burdwanensis, O. translineatus, and O. wangi represent a cluster of very similar species. Finally, the last grouping in Table
The degree of fusion of the metapodal platelet with the various surrounding shields may vary significantly intraspecifically, as seen in O. mooiensis (incl. syn. O. gamagarensis;
Intraspecific variation in shield fusions may occur in males too. For instance, some males that we identified as O. placidus have the peritrematal shield fused to the hologastric shield, just like the male of O. venetus, and others have the peritrematal shield free posteriorly, like that of the male of O. sellnicki (
At present, the chaetotaxy and the ornamentation of the dorsal shield are not clearly described for most Ololaelaps species (Table
Presently, differences in dimensions of the dorsal, sternal, and hologastric shields are only useful to separate species with marked differences, i.e., with elongate (e.g., O. tasmanicus) vs broad shields (e.g., O. placentula), because intraspecific variation is not sufficiently known. Ratios of length/width could be particularly useful, but they also vary intraspecifically, e.g., the sternal shield of O. venetus appears to have a length/width ratio of 0.8–1.0 (
The position of seta st4 and poroid iv3 is difficult to use as a diagnostic character because it is not easy to determine whether they are on the shield margin, on the adjacent soft cuticle, or on the endopodal plate. This body region being the point of meeting of three shields (sternal, endopodal, hologastric) renders its study more difficult, obscuring the position of st4 and iv3, especially if they are inserted on soft cuticle, which can be folded above or underneath shields’ margins. Examining several specimens for each species can help, as well as making observations at different focal depths. We suspect that in most cases where st4 (and iv3) appears on the endopodal plate (e.g., O. burdwanensis, O. sitalaensis, O. translineatus), it is actually inserted on soft cuticle that overlaps the plate. Note that the position of st4 and iv3 are relatively stable within genera or even families of Gamasina, whether on soft cuticle, on metasternal platelets or (more rarely) on the sternal shield (e.g.,
The ornamentation of the hologastric shield shows species-specific patterns, such as inverted V or U-shaped ridges in O. formidabilis and undescribed species, as well as the shape of cells in the reticulation pattern (e.g.,
Our knowledge of the gnathosoma of Ololaelaps indicates limited variation between species. For instance, the internal malae have two pairs of projections in the females of all species where the hypostome has been described (O. caucasicus, dililoensis, formidabilis, mooiensis, placentula, placidus, sellnicki, ussuriensis, venetus, wangi) except for O. sitalaensis which lacks the lateral pair, based on the illustration in
Idiosomal adenotaxy differs between laelapid species (
While the legs of Ololaelaps species mostly bear simple and slender setae, there is interspecific variation in the shape of setae. This should be investigated and exploited for species diagnostics (see examples in the genus description above).
We sincerely thank Evert Lindquist, Victoria Nowell (Agriculture and Agri-Food Canada, AAFC), Bruce Halliday (CSIRO) and Alireza Nemati (Shahrekord University, Iran) for valuable comments on the manuscript; Roberto Nannelli (Centro di Ricerca per l’Agrobiologia e la Pedologia, Firenze, Italy) for examining and photographing the type material of O. formidabilis in the Berlese Collection; Hans Klompen (Ohio State University), Jose Fernández-Triana, Scott Brooks, Jean-François Landry, Serge Laplante, Aleš Smetana and Monty Wood (AAFC) for useful advice on species boundaries or nomenclature; Tanya Durr (AAFC) for assistance in finding references, and Bruce Halliday (CSIRO), Gilberto de Moraes (University of São Paulo, Brazil), Diana Rueda (University of São Paulo) and Axel Christian (Senckenberg Museum of Natural History Görlitz) for sharing literature with us; Vasily Grebennikov (Canadian Food Inspection Agency) and Jessica Hsiung (AAFC) for translating parts of Russian and Chinese papers; the State Ministry of Research and Technology of Indonesia (RISTEK) for the research permit; the Indonesian Institute of Science (LIPI) and the Ministry of Forestry (PHKA) for the collecting permit; the village heads and local site owners for granting access to their properties, and the many colleagues and helpers for support in the field. Financial support was provided by the German Research Foundation (DFG) in the framework of the collaborative German – Indonesian research project CRC990 (EFForTS).