Corresponding author: Thorsten Assmann (
Academic editor: J. Liebherr
The carabids of the tribe
Assmann T, Boutaud E, Buse J, Drees C, Friedman A-L-L, Harry I, Khoury F, Orbach E, Renan I, Schmidt C, Schmidt K, Wrase DW, Zumstein P (2021) The ground beetle tribe Platynini Bonelli, 1810 (Coleoptera, Carabidae) in the southern Levant: dichotomous and interactive identification tools, ecological traits, and distribution. In: Spence J, Casale A, Assmann T, Liebherr J, Penev L (Eds) Systematic Zoology and Biodiversity Science: A tribute to Terry Erwin (1940–2020). ZooKeys 1044: 449–478.
The
In order to support ecologists, environmentalists, and conservationists, but also citizen scientists, we have developed identification keys for this tribe. We present these taxonomic tools in two forms: a “classical” dichotomous key and an interactive key with further identification tools like character matrices.
In the pioneer phase of entomological exploration of the southern Levant, some species of
This study is based on the examination of specimens (i) observed and/or collected during field trips of the authors in Israel, Jordan and Egypt (between 2004 and 2020), (ii) collected in context of ecological surveys (e.g.,
Abbreviations of collections (used also in Suppl. material
The basic approach to identify ecological and distributional traits and threats follows our previous work (
We prepared the photos with the microscope and camera mentioned in the previously cited publications (
We created an interactive key in the Xper3 version 1.5.2, a collaborative web-based platform (
As “items”, the species are incorporated into the database (“
Altogether, we could verify more than 800 specimens of platynines from the southern Levant. We can present here first faunistic records for countries of the southern Levant:
Following the database and our literature survey, a total of 13 species occurs in the southern Levant, and one additional species may occur in Northwest Syria. In the following key, we list the latter species in parentheses. We incorporated the 14 species in both identification keys, the classic dichotomous identification key and the interactive key.
Within the
Medium sized species of slender and flat habitus, often with noticeable metallic luster. Head with two pairs of supraorbital setae. Mandible scrobe (groove on the basal half of the mandible laterally) without a seta. Antennae pubescent from the 2nd, 3rd, or 4th antennomeres. Penultimate labial palpomere with two setae on its anterior margin. Process of prosternum not margined apically. Protibia apically with deep emargination (antenna cleaner) not enlarged. Tarsi on the upper side rarely pubescent. Ventral edges of claws smooth. Both parameres of the median lobe of aedeagus spatulate and ovate. In addition, Figs
1 | Upper side with numerous hairs, especially on pronotum and elytra |
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– | Upper side without hairs, only regular setae (supraorbital setae, series umbilicata, etc.) |
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2 | Pronotum cordiform, almost as long as wide (Fig. |
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– | Pronotum clearly wider than long (Fig. |
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3 | Labrum clearly convex; apical margin rounded; mandibles shorter, ca. 2× as long as labrum (Fig. |
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– | Labrum flat or slightly convex; apical margin almost straight (sometimes slightly convex or concave); mandibles longer, in most species > 2.5 × as long as labrum (Figs |
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4 | Pronotum wider, ca. 1.3× wider than long; lateral bead broader, punctuation stronger (Fig. |
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– | Pronotum longer, ca. 1.2× wider than long; lateral bead slender, punctuation weaker (Fig. |
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5 | Hind angles of pronotum ca. rectangular, sometimes produced in sharp denticle, sometimes bases of pronotum somewhat convex. Upper side colorful either with blue to violet metallic luster or head and pronotum with green or blue metallic luster and elytra matt brown with darkening in central and apical part (Figs |
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– | Hind angles of pronotum entirely rounded or obtuse, sometimes slightly produced into minute denticle (Figs |
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6 | Head and pronotum with green or bluish green metallic luster, elytra matt brown with darkening in central and apical part. Hind angles of pronotum without seta (Fig. |
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– | Head, pronotum and elytra with blue to violet luster (Figs |
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7 | First antennomere in apical part enlarged, behind a small constriction (“trumpet-shaped”). Pronotum 1.26–1.38 wider than long (Fig. |
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– | First antennomere regularly shaped, in apical part neither with a constriction nor widened at apex. Pronotum 1.16–1.27 wider than long (Fig. |
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8 | Upper side green and/or coppery; with broad yellow elytral margin; yellow coloration on pronotal margin narrower (Fig. |
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– | Upper side variable, from black and brown to vivid metallic luster, but without yellow margin on elytra or pronotum (Figs |
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9 | Third elytral intervals with (5-) 6 (-7) setiferous punctures. Upper side with metallic luster, mostly elytra green and forebody (head, pronotum) reddish or coppery, rarely black (Fig. |
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– | Third elytral intervals with (2-) 3 (-4) setiferous punctures. Upper side of different color |
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10 | Elytra brownish, femora reddish to pale brown; tibiae darker. Apex of 3rd antennal segment with additional (sometimes only few) small hairs beside regular erect setae (Fig. |
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– | Elytra with metallic luster or black or dark brown to black. Apex of 3rd antennal segment only with regular setae |
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11 | Larger species (> 8.5 mm) with small and wide pronotum (mostly more than 1.3× wider than long). Microsculpture of elytra isodiametric and strongly developed, also in males. Vivid metallic green or bronze, rarely black. With depression in 5th stria and adjacent intervals of apical third of elytra (Fig. |
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– | Smaller species up to 9.0 mm with larger and slender pronotum (mostly < 1.3 × wider than long). Microsculture of elytra isodiametric, but less developed. Never vividly metallic colored, elytra dark brown to black, completely without or with slight metallic luster | 12 |
12 | Pronotum wider, 1.2 to 1.3× wider than long, lateral margin continuously rounded between maximum width and hind angles (Fig. |
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– | Pronotum narrower, < 1.2× wider than long, lateral margin from maximum width towards hind angles slightly straight or slightly concave (Figs |
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13 | Pronotum 1.1 to 1.2× wider than long, lateral margin from maximum width towards hind angles straight or very slightly concave, hind angles less rounded, fore angles pronounced (Fig. |
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– | Pronotum ≤ 1.1× wider than long, lateral margin from maximum width towards hind angles straight, hind angles more rounded, fore angles stronger rounded (Fig. |
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The dichotomous identification key has 13 alternatives. The number of alternatives or clusters of alternatives, respectively, that have to be considered for the identification of a single species ranges from two to nine.
Pronotum of
The interactive key is based on 16 characters with the number of character states ranging from 2 to 12 (Table
The states for the characters punctuation of pronotum and the number of setiferous punctures in the 3rd elytral interval are not available for all species. In these cases, additional photographs would have been necessary to make the hard-to-recognize character states of these species clear to the users of the key. The number of setiferous punctures in the 3rd elytra interval is difficult to detect in strongly pubescent species; and the intensity of the punctation of the pronotum can be inferred by a user only by specimen comparisons or with Figs
Pronotum of
Apical margins of elytra of
Overview of the characters, number of their states, and number of species for which the character states are applicable in the Xper3 database “
Character (“descriptor“) | Number of character states (categories) | Number of species for which character states are applicable |
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Body length (numerical) | Variable | 14 |
Coloration of upper side (head, pronotum, elytra) (categorical) | 13 | 14 |
Metallic luster of upper side (categorical) | 3 | 14 |
Coloration of legs (categorical) | 4 | 14 |
First antennomere (categorical) | 2 | 14 |
Labrum (categorical) | 2 | 14 |
Mandibles (categorical) | 2 | 14 |
Pronotum hind angles (categorical) | 2 | 14 |
Pronotum lateral margin (categorical) | 3 | 14 |
Pronotum proportion (categorical) | 4 | 14 |
Pronotum punctation (categorical) | 2 | 3 |
Hairs on upper side (pronotum, elytra) (categorical) | 2 | 14 |
3rd elytral interval, number of setiferous punctures (numerical) | Variable | 12 |
5th elytral stria and adjacent intervals (categorical) | 2 | 14 |
Apical margin of elytra (categorical) | 2 | 14 |
Without the option for applying these character states in some species, the interactive key helps to identify the species more quickly than the dichotomous one: If the character with the most states (esp. coloration of upper side) is included for the first decision, only 1 to 3 decisions are necessary for the identification of a species. However, if characters with a low number of states are prioritized, the number of decisions increases but remain lower than for the dichotomous identification key and ranges from 2 to 5.
Our Xper3 knowledge data base “
After a discussion with the two laypersons who tested several keys generated by Xper3, we weighted four characters strongly, two weakly, and all others medium. By using the options ‘use weights’ for the characters and ‘prioritize characters with fewer states’ we generated a single access key which has 17 alternatives (Suppl. material
The interactive identification key is available at
The database is available at
The sdd file (in xml format) with the information incorporated in the knowledge database ‘
Apical part of left elytron with 5th stria and adjacent intervals of
Apical part of Head of
Fully winged and flight-active; able to quickly colonize restored habitats (
Stream banks and margins of winter ponds and artificial water reservoirs where vegetation is mostly sparse (Fig.
Spring breeder (
Throughout Europe (without North Scandinavia), southwards to North Africa, eastwards to Cyprus, Asia Minor, and southern Levant (
Already listed by
In most parts of the distribution range abundant, not threatened (e.g.,
Fully winged (n = 2). No flight observation known.
We know the habitat only from one site in Iran: partly overgrown gravel banks along streams at ca. 2350 m a.s.l. Also in Kyrgyzstan at high altitude (2000 m).
Unknown.
From Georgia to Central Asia and Pakistan (
For their description of the species,
Habitats of
Fully winged and flight-active (pers. obs.).
Known only from winter ponds (Fig.
Spring breeder, adults hibernate (pers. obs.).
The species is distributed from Southwest Europe to Central Asia, southwards to Israel and Iran (
In the southern Levant recorded exclusively from four sites in Israel, three of them from winter pools in the Central Coastal Plain (Berekhat Ya’ar (sometimes also written Breikhat Ya’ar), Dora Pool and Ga’ash Pool; from the latter two sites only single specimens). A large population exists also in the swamps north of the Sea of Galilee.
In contrast to
Fully winged and flight-active (
A hygrophilous species in a wide range of habitats: from freshwater to brackish water habitats like winter ponds, shaded and unshaded streams, lakes, swamps, etc. Very abundant in the swamps north of the Sea of Galilee (Figs
Spring breeder (reproduction. March to May), adults hibernate.
From western Europe to Central Asia, northwards to Central Europe, southwards to North Africa, eastwards to Central Asia (
From the Mediterranean climate zone to desert zone (Central Negev and Wadi Hasa). Egypt (Sinai, several sites:
We did not find the species in the extremely arid and hot areas around the Dead Sea or at the springs in the Arava Valley. Also in the light catches from the En Gedi Nature Reserve (kept in the
In contrast to some other
Fully winged.
Winter ponds and small associated streams. The water reservoir of Merom Golan hosts also a population (Fig.
Spring breeder.
From Southeast Europe to Iran, not in North Africa (
North Israel (Upper Galilee, Hermon, Golan Heights).
Fully winged and flight active.
Winter ponds and small associated streams. The water reservoir of Merom Golan hosts also a population (Fig.
Spring breeder, adults hibernate.
From Southeast Europe to Asia Minor and southern Levant (
Known only from Golan Heights.
Fully winged and flight active (pers. obs.). A species with an abrupt northward distributional extension in Central Europe (
Wetlands, mostly winter ponds, rarely on riverbanks of streams or in semi-open floodplain woodlands (e.g., swamps with
Spring breeder, tenerals at least sometimes also in March; adults hibernate.
The nominate subspecies from Southwest Europe to West Siberia and southern Levant (including Sinai,
In the Mediterranean climate zone (Lebanon, North Israel, Egypt: North Sinai) (
Fully winged and flight active.
Riparian species of permanent and temporary streams, both sun-exposed and shaded, e.g., by
At type locality a spring breeder with copulation in March and April, in May already tenerals; adults hibernate (pers. obs.).
Endemic in a small range of the southern Levant (
Few localities from North Lebanon to the West Golan Heights (
Due to excessive water withdrawal, streams in Israel and Lebanon are drying out faster, and their flood dynamics are changing. This results in a deterioration of the gravel banks, which largely become overgrown and are then no longer available as habitat for the species. Headwater areas are developed into recreational areas and naturally occurring microhabitats are destroyed (e.g., Banyas Waterfalls). Threatened due to the small range and poor conservation status of flowing waters in the southern Levant.
Fully winged.
Unknown.
Unknown; types are from May and June (
Endemic in South and central East Turkey, probably also in Northwest Syria (
Unknown (
Fully winged, apical wing part of variable size (
In Europe a widespread species of open habitats, esp. of grassland (meadows, pastures) and arable fields (
Spring breeder; adults hibernate (
From West Europe to Central Asia, southwards to North Africa, northwards to Scandinavia. The nominate subspecies in Turkey, Syria, and Cyprus (
Mainly in the Mediterranean climate zone of the southern Levant, but a few specimens were collected in the Dead Sea Area (Nahal Qumeran, ‘Enot Zuqim, 1993) and in the southern mountains of Sinai (Jebel Katarina, Wadi Feiran, Wadi Wattir, 1971–1976).
Hind wings are longer than the elytra; we know of a flight observation from a car net (Michael Schuelke, personal observation).
Softwood floodplain woodlands with
All our specimens from Israel are from March to May; probably a spring breeder with summer larvae.
From southwest Europe (
First record from Tanur (Tanur Waterfalls, Nahal ‘Iyyon Nature Reserve) (
The damages to water bodies of streams (exploitation, pollution, reduced dynamics of floods due to drainage) (
Fully winged (n = 4) and flight activity (at light, own observation from Spain).
In open, sparsely vegetated habitats on loamy or sandy soil, also in clearings of forests. Apparently not restricted to wetlands, even if the species is found there from time to time. (
Unknown.
From southwest Europe (Spain) to Asia minor and southern Levant, including Cyprus, southwards to North Africa (
In the Mediterranean part of Israel (Upper Galilee, Lower Galilee, Golan Heights, Carmel Ridge, Central coastal Plain, Judean Hills). Already mentioned by
The tip of the median lobe of the aedeagus shows remarkable variation across the distribution range, but the endophallus seems to be of the same shape.
Fully winged (n = 8) and probably flight-active.
Batha (with
Probably an autumn and winter breeder.
From southwestern Europe to Asia Minor and southern Levant (
Few records from Israel (
Fully winged (n = 4).
In Pakistan, in thorn scrub forests (
In Saudi Arabia, from November to May (
From Egypt to Iran and Pakistan (
Known from Egypt, incl. Sinai (
Computer- and/or Internet-based identification options are increasing. Many of them work very well including, for example, some applications for smartphones, tablets, or personal computers. Some of these applications are based on machine learning (esp. convolutional neural network, CNN) (cf.
The keys presented here have different numbers of decisions up to species identification. Especially the interactive key allows a fast identification. However, one character, the coloration of the upper side, has many states. In case of polychromatic species, the coloration can only be used to unambiguously identify some individuals of a species. This applies for example to the very variable species
We do not know whether the identification of ground beetle species with the help of an interactive key is as reliable as with a classical dichotomous key. Up to now, we had the impression that an identification is more reliable if a combination of character states leads to the actual result as typically encounter in classical dichotomous identification key. Often, features such as coloration or body length can validate an identification. If one would like to have this additional information with the use of the interactive key, one should consult additional tools of our Xper3 knowledge base. These include the information that can be found in the species (“items”) section under “Details” and photos under “Pictures”. In addition, under the tools there is also the “Description Matrix” with the character states for each species and character.
Interactive and single access keys, created by Xper3, seem to have a greater efficiency than classical dichotomous keys, not only in our study, but also for coccinellids in France (
We see a great advantage of knowledge bases created on the Xper3 platform in the possibility to remain dynamic. This implies that a knowledge base could be updated when new knowledge is available, e.g., by adding further taxa. In the case of the knowledge base ‘
At least eight species of
The reproductive period of the two
A complex system with different photoperiodic control of previtellogenesis and vitellogenesis (and probably also spermatogenesis) determines the reproductive rhythmicity of many ground beetles (
Many
Continued water withdrawal and groundwater lowering are severely threatening the stream habitats in the Levant (
The hygrophilous
We dedicate this work to the late Terry Erwin for his entomological performance and enthusiasm for carabidology. With this contribution, we would like to thank him for joint excursions and his manifold support for us. His splendid life’s work in carabidology and biodiversity research stimulates us greatly and we will never forget him.
We thank Joachim Schmidt (Rostock) for his helpfulness in questions concerning the determination and synonymy of platynines. Christoph Reuter (Berlin) very generously supported us with carabid specimens from Lebanon and other countries in Southwest Asia. The same applies to Peer H. Schnitter (Halle), Kamil Orszulik (Frýdec-Místek), and Zbynek Kejval (Domažlice). Special thanks go to James Liebherr (Ithaca) and Riccardo Sciaky (Milan) for valuable comments on the manuscript during the review process. We thank also the Israel Nature Protection Authority for giving permits to study insects in the nature reserves. Thanks goes also to those scientists who supported us both in the field and in the collections, esp. Tamar Dayan, Netta Dorchin, the late Amnon Friedberg (all
Material examined: Records of the
table
Single access identification key generated by Xper3, using weights of the characters and prioritization of characters with few states
Hypertext Markup Language (html)
Exported knowledge database ‘
Extensible Markup Language (xml)